Mutations in mads-box genes and uses thereof

ABSTRACT

Aspects of the disclosure relate to plants, such as Solanaceae plants containing one or more of a mutant Solyc04g005320 gene (or a homolog thereof), a mutant Solycl2g038510 gene (or a homolog thereof), and a mutant Solyc03gl14840 gene (or a homolog thereof), as well as methods of producing such plants. In some aspects, such plants have one or more improved traits, such as modified inflorescence architecture, modified flower number, modified fruit number, higher yield, higher quality products, and higher fruit productivity.

CROSS-REFERENCE TO RELATED APPLICATIONS

This application claims the benefit of the filing date of U.S.Provisional Application No. 62/507,369, filed on May 17, 2017. Theentire contents of this referenced application are incorporated byreference herein.

GOVERNMENT SUPPORT

This invention was made with government support under IOS-1523423 andIOS-1237880 awarded by the National Science Foundation. The governmenthas certain rights in the invention.

BACKGROUND

The architectures of plant reproductive shoot systems—inflorescences—aremajor determinants of crop yield, and modified inflorescence complexitywas a recurring target during crop domestication and improvement(Doebley et al., 2006; Meyer and Purugganan, 2013). Prominent examplesinclude the cereal crops barley, maize, rice and wheat, for which humansselected variants with greater branching to increase flower and grainproduction (Ashikari et al., 2005; Boden et al., 2015; Doebley et al.,1997; Huang et al., 2009; Jiao et al., 2010; Komatsuda et al., 2007;Ramsay et al., 2011). Yet, for many crops, particularly fruit-bearingspecies such as grape and tomato, inflorescence architecture has changedlittle from wild ancestors, and therefore has been underexploited inbreeding (Lippman et al., 2008; Mullins et al., 1992; Peralta andSpooner, 2005).

SUMMARY

Aspects of the present disclosure relate to compositions, such as novelgenetic variants of plants, and methods for generating the compositions,which have favorable traits, such as yield-related traits. In someaspects, the combination of mutations in the novel genetic variantsincrease inflorescence and fruit production. In other aspects, mutationsin one or more of the genes of the genetic variants can be used tocreate a quantitative range of inflorescence types, such as thedevelopment of weakly branched genetic variants that results in higherflower and fruit production.

In some aspects, the disclosure provides a genetically-alteredSolanaceae plant (e.g., a tomato plant) comprising a mutantSolyc04g005320 gene or a homolog thereof. In some embodiments, themutant Solyc04g005320 gene or homolog thereof is a null allele or ahypomorphic allele. In some embodiments, the genetically-alteredSolanaceae plant (e.g., tomato plant) is heterozygous or homozygous forthe mutant Solyc04g005320 gene or homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) further comprises a mutant Solyc12g038510 gene or ahomolog thereof, a mutant Solyc03g114840 gene or a homolog thereof, orboth a mutant Solyc12g038510 gene or a homolog thereof and a mutantSolyc03g114840 gene or a homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) further comprises a mutant Solyc12g038510 gene or homologthereof and the mutant Solyc12g038510 gene or homolog thereof is a nullallele or a hypomorphic allele. In some embodiments, thegenetically-altered Solanaceae plant is heterozygous or homozygous forthe mutant Solyc12g038510 gene or homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) further comprises a mutant Solyc03g114840 gene or ahomolog thereof and the mutant Solyc03g114840 gene or homolog thereof isa null allele or a hypomorphic allele. In some embodiments, thegenetically-altered Solanaceae plant is heterozygous or homozygous forthe mutant Solyc03g114840 gene or homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) further comprises both a mutant Solyc12g038510 gene or ahomolog thereof and a mutant Solyc03g114840 gene or a homolog thereof,each of which are independently a null allele or a hypomorphic allele.In some embodiments, the genetically-altered Solanaceae plant isheterozygous or homozygous for the mutant Solyc12g038510 gene or homologthereof and is heterozygous or homozygous for the mutant Solyc03g114840gene or homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) comprises the mutant Solyc04g005320 gene or homologthereof, the mutant Solyc12g038510 gene or homolog thereof, and themutant Solyc03g114840 gene or homolog thereof, and wherein each is ahypomorphic allele. In some embodiments, the genetically-alteredSolanaceae plant (e.g., tomato plant) is heterozygous or homozygous forthe mutant Solyc04g005320 gene or homolog thereof, is heterozygous orhomozygous for the mutant Solyc03g114840 gene or homolog thereof and isheterozygous or homozygous for the mutant Solyc03g114840 gene or homologthereof.

In some embodiments, the mutant Solyc04g005320 gene or homolog thereofis a hypermorphic allele. In some embodiments, the genetically-alteredSolanaceae plant (e.g., tomato plant) is heterozygous or homozygous forthe mutant Solyc04g005320 gene or homolog thereof.

In some embodiments, the genetically-altered Solanaceae plant (e.g.,tomato plant) further comprises a mutant Solyc12g038510 gene or ahomolog thereof, a mutant Solyc03g114840 gene or a homolog thereof, orboth the mutant Solyc12g038510 gene or homolog thereof and the mutantSolyc03g114840 gene or homolog thereof.

In other aspects, the disclosure provides a genetically-alteredSolanaceae plant (e.g., a tomato plant), comprising a mutantSolyc12g038510 gene or a homolog thereof and a mutant Solyc03g114840gene or a homolog thereof, wherein the genetically-altered Solanaceaeplant is homozygous for the mutant Solyc12g038510 gene or homologthereof and heterozygous for the mutant Solyc03g114840 gene or homologthereof. In some embodiments, the mutant Solyc12g038510 gene or homologthereof is a null allele or a hypomorphic allele and the mutantSolyc03g114840 gene or homolog thereof is a null allele or a hypomorphicallele.

In some embodiments of any one of the genetically-altered Solanaceaeplants (e.g., a tomato plant) provided herein, the mutant Solyc04g005320gene or homolog thereof, the mutant Solyc12g038510 gene or homologthereof, and/or the mutant Solyc03g114840 gene or homolog thereof isintroduced by technical means. In some embodiments of any one of thegenetically-altered Solanaceae plants (e.g., a tomato plant) providedherein, the mutant Solyc04g005320 gene or homolog thereof, the mutantSolyc12g038510 gene or homolog thereof, and/or the mutant Solyc03g114840gene or homolog thereof is introduced by chemical or physical means. Insome embodiments of any one of the genetically-altered Solanaceae plants(e.g., a tomato plant) provided herein, the mutant Solyc04g005320 geneor homolog thereof, the mutant Solyc12g038510 gene or homolog thereof,and/or the mutant Solyc03g114840 gene or homolog thereof is introducedusing CRISPR/Cas9, chemical mutagenesis, radiation,Agrobacterium-mediated recombination, viral-vector mediatedrecombination, or transposon mutagenesis. In some embodiments of any oneof the genetically-altered Solanaceae plants (e.g., a tomato plant)provided herein, the plants are provided with the provision that plantsexclusively obtained by means of an essentially biological process areexcluded.

In other aspects, the disclosure provides a crop harvested from agenetically-altered Solanaceae plant (e.g., a tomato plant) of any oneof the above embodiments or of any other embodiment described herein.

In yet other aspects, the disclosure provides a seed for producing agenetically-altered Solanaceae plant (e.g., a tomato plant) of any oneof the above embodiments or of any other embodiment described herein.

In other aspects, the disclosure provides a method for producing agenetically-altered Solanaceae plant (e.g., a tomato plant), the methodcomprising introducing a mutation into a Solyc04g005320 gene or ahomolog thereof in a Solanaceae plant, thereby producing agenetically-altered Solanaceae plant containing a mutant Solyc04g005320gene or homolog thereof. In some embodiments, the mutation is introducedusing CRISPR/Cas9. In some embodiments, the mutation produces a nullallele or a hypomorphic allele of the Solyc04g005320 gene or homologthereof.

In some embodiments of any one of the methods provided herein, themethod further comprises introducing into the Solanaceae plant amutation into a Solyc12g038510 gene or a homolog thereof, introducing amutation into a Solyc03g114840 gene or a homolog thereof, or introducingthe mutation into the Solyc12g038510 gene or homolog thereof andintroducing the mutation into the Solyc03g114840 gene or homologthereof. In some embodiments, the mutation(s) is/are introduced usingCRISPR/Cas9.

In some embodiments of any one of the methods provided herein, thegenetically-altered Solanaceae plant (e.g., a tomato plant) containingthe mutant Solyc04g005320 gene or homolog thereof is crossed to anothergenetically-altered Solanaceae plant (e.g., a tomato plant) comprising amutant Solyc12g038510 gene or homolog thereof, a mutant Solyc03g114840gene or homolog thereof, or both the mutant Solyc12g038510 gene orhomolog thereof and the mutant Solyc03g114840 gene or homolog thereof,thereby producing a genetically-altered Solanaceae plant (e.g., a tomatoplant) containing the mutant Solyc04g005320 gene or homolog thereof andthe mutant Solyc12g038510 gene or homolog thereof, the mutantSolyc03g114840 gene or homolog thereof, or both the mutantSolyc12g038510 gene or homolog thereof and the mutant Solyc03g114840gene or homolog thereof.

In other aspects, the disclosure provides a genetically-alteredSolanaceae plant (e.g., a tomato plant) produced or obtainable by amethod of any one of the above embodiments or of any other embodimentdescribed herein.

BRIEF DESCRIPTION OF THE DRAWINGS

The following drawings form part of the present specification and areincluded to further demonstrate certain aspects of the presentdisclosure, which can be better understood by reference to one or moreof these drawings in combination with the detailed description ofspecific embodiments presented herein. It is to be understood that thedata illustrated in the drawings in no way limit the scope of thedisclosure.

FIGS. 1A-1K show the s2 inflorescence architecture variant branches dueto delayed meristem maturation. FIG. 1A shows a typical wild type (WT)tomato plant with unbranched, multi-flowered inflorescences and jointedpedicels (dotted asterisk in inset). Numbers in FIGS. 1A-1C indicateflowers per inflorescence (mean±SEM, N=number of inflorescences).Striped arrowheads indicate successive inflorescences. P: two-tailed,two-sample t-test compared to WT. FIG. 1B shows the highly branchedinflorescences and jointed pedicels of s mutants. White arrowheadsindicate branch points. FIG. 1C shows the s2 mutant with moderatelybranched inflorescences and jointless pedicels (asterisk). FIG. 1D showsquantification of inflorescence branching events in WT, s, and s2. FIG.1E shows phenotypic classes in a WT×s2 F2 population. The segregationratio for the jointless pedicel phenotype and the branched inflorescencephenotype (s2) is given. Asterisks mark jointless pedicels. Scale barsin FIGS. 1A-1C and lE=1 cm. FIGS. 1F-1H show the transition meristem(TM), sympodial inflorescence meristem (SIM), and floral meristem (FM)from WT (FIG. 1F), s (FIG. 1G), and s2 (FIG. 1H). Scale bars in FIGS.1F-1H represent 100 μm. L, leaf. F, flower. Schematics depict developinginflorescences. Lines, internodes; circles, FMs/flowers; arrowheads,SIMs. Overproliferating branches are indicated in bolded line. FIG. 1Ishows PCA of 2,582 dynamically expressed genes in the vegetativemeristem (VM), TM, SIM, and FM of WT, s, and s2, determined by RNA-seq.FIGS. 1J-1K show expression (z-score normalized) of TM (FIG. 1J) and FM(FIG. 1K) marker genes in the vegetative (VM) meristem, TM and FM stageof meristem maturation of WT and mutant (s and s2). Cluster of geneswith moderately (left) and strongly (right) delayed expression patternare shown. Dashed lines indicate median expression with dot-filled-inarea representing the 5^(th) and 95^(th) quantile.

FIGS. 2A-2N show that mutations in two SEPALLATA MADS-box genes cause s2branching. FIG. 2A shows mapping-by-sequencing of s2. Ratio ofSNP-ratios (s2/M82) between different pools of segregating phenotypicclasses (top: s2/WT; middle: s2/j2; bottom: j2/WT) is shown forchromosome 3 and 12. FIG. 2B shows the j2 mapping interval includes theSEP4 homolog Solyc12g038510. FIG. 2C shows Genomic Illumina-sequencereads showing a breakpoint in Solyc12g038510 (left), and PCR showing aCopia/Rider transposon insertion in the first intron of Solyc12g038510in s2 mutants (right). The sequence corresponds to SEQ ID NO: 89. FIG.2D shows Sashimi plots of normalized RNA-seq reads (reads per million,RPM) for Solyc12g038510 in WT (top) and s2 (bottom) floral meristems. Anintronic transcriptional start site leads to out-of-frame Solyc12g038510transcripts in s2 mutants. Numbers indicate reads supportingsplice-junctions and alternative splicing in s2 is highlighted in thebottom panel by diagonal line filling. FIG. 2E shows the generation ofj2^(CR) null mutations by CRISPR/Cas9 using two single-guide RNAs(sgRNA, target1 and target2; arrows). Black arrows indicate forward (F)and reverse (R) primers used for genotyping and sequencing. Sequences ofj2^(CR) allele 1 (a1) and a2 are shown. sgRNA targets andprotospacer-adjacent motif (PAM) are indicated in bold font anddeletions by dashes. Insertions are indicated in italic font andsequence gap length is shown in parentheses. From top to bottom,sequences correspond to SEQ ID NOs: 90-92. FIG. 2F shows inflorescencesand fruits from WT and j2^(CR) mutants showing unbranched inflorescencewith jointless pedicels for j2^(CR). White and dotted asterisks indicatejointed and jointless pedicels, respectively. FIG. 2G shows acomplementation test between j2^(CR) and j2^(TE) (jointless pedicels;asterisks). FIG. 2H shows that the ej2 mapping interval includes theSEP4 homolog Solyc03g114840. FIG. 2I shows the Genomic Illumina-sequencereads showing a breakpoint in Solyc03g114840 and PCR revealing a 564 bpinsertion in the 5th intron of Solyc03g114840 in s2 mutants. Thesequence corresponds to SEQ ID NO: 93. FIG. 2J shows Sashimi plots forSolyc03g114840 RNA-seq reads in WT and s2 floral meristems indicatingpartial exon skipping and intron retention in s2 mutants. FIG. 2K showsthe generation of ej2^(CR) null mutations by CRISPR/Cas9. From top tobottom, sequences correspond to SEQ ID NOs: 94-97. FIG. 2L showsunbranched ej2^(CR) mutant inflorescences with extremely long sepals(arrowheads) and pear-shaped fruits. Scale bars=1 cm. FIG. 2M showsunopened flowers showing the weak natural ej2^(w) allele causes longersepals and fails to complement ej2^(CR). FIG. 2N shows quantification ofrelative sepal length (sepal length/petal length f SEM, N=number offlowers) for genotypes in FIG. 2M. P: two-tailed, two-sample t-testcompared to WT.

FIGS. 3A-3F show the ej2^(w) variant arose during domestication and wasselected during breeding of large-fruited cultivars. FIG. 3A showsdistribution of the ej2^(w) allele in wild tomato species, earlydomesticates (landraces, S. lyc. var. cerasiforme), and cultivars (S.lycopersicum)(N=number of accessions). FIG. 3B shows relative sepallength (sepal length/petal length) from a subset of accessions in FIG.3A homozygous EJ2 and ej2^(w). FIG. 3C shows relative sepal length in asubset of confirmed landraces (Blanca et al., 2015). FIG. 3D shows PCRgenotyping for the ej2^(w) allele in 10 landraces with the longest andshortest sepals. S. pimpinellifolium (S. pim) was used as a WT control.FIG. 3E shows inflorescences and flowers (inset) of the accessions withthe shortest and longest sepals. See asterisks in FIG. 3D. Numbersindicate relative sepal length. FIG. 3F shows PCR genotyping in 258cultivars shows enrichment of the ej2^(w) allele in large-fruited types.Data in FIGS. 3B, 3C, and 3E are means (±SEM, n=10 flowers peraccession). N=number of accessions. P: two-tailed, two-sample t-test.Scale bars=1 cm.

FIGS. 4A-4D show that breeders overcame negative epistasis between j2and ej2 by selecting suppressors of s2 branching in elite germplasm.FIG. 4A shows PCR genotyping of 153 elite breeding lines for j2^(TE) andej2^(w) reveals the jointless germplasm is dominated by the j2transposon allele and contains many j2^(TE) ej2^(w) double mutants.Number of accessions is indicated in parenthesis. FIG. 4B shows PCRgenotyping of 31 jointless inbreds and hybrids from 4 major seedcompanies for ej2^(w). Asterisks indicate j2^(TE) ej2^(w) doublemutants. FIG. 4C shows representative images of phenotypic classes foundinj2^(TE) ej2^(w) double mutants isolated from an S. pimpinellifolium×s2F2 population. N indicates number of plants and percentage of plants ineach phenotypic class is indicated in parentheses. FIG. 4D showsmapping-by-sequencing a suppressor of s2 to a 3 Mbp interval onchromosome 2 containing 167 genes. DNA from pools of s2 and suppresseds2 plants was sequenced and the ratio (suppressed s2/s2) of theSNP-ratios (S.pin/s2) is presented.

FIGS. 5A-5I show that redundancy among three SEP4 genes regulatesinflorescence branching and flower development. FIG. 5A shows thephylogenetic tree of SEP proteins in Arabidopsis and tomato. Bootstrapvalues (%) for 1000 replicates are shown. FIG. 5B shows normalized geneexpression (RPKM) for TM5 and TM29 (left) and the SEP4 sub-clade (right)during meristem maturation (VM, vegetative meristem; TM, transitionmeristem; FM, floral meristem; SIM, sympodial inflorescence meristem;SYM, sympodial shoot meristem). FIG. 5C shows yeast two-hybrid assaysshowing heteromeric interactions for Solyc04g005320, J2, and EJ2, andhomomeric interactions for Solyc04g005320 and J2 (3-AT,3-amino-1,2,4-triazole; L, leucine; T, tryptophan; H, histidine; e.v.,empty vector). FIG. 5D shows the summary of results in FIG. 5C; (−) nointeraction; (+) interaction; (++) strong interaction. FIG. 5E shows thelonger inflorescence of a Solyc04g005320^(CR) mutant (hereafter referredto as long inflorescence^(CR); lin^(CR)) compared to WT. Numbersindicate flowers per inflorescence (mean±SEM, N=10 inflorescences). P:two-tailed, two-sample t-test. Scale bar=1 cm. FIG. 5F shows the longerinflorescence of a Solyc04g005320^(CR) mutant in S. pimpineiolium (S.pim lin^(CR)) compared to S. pimpinellifolium WT. FIG. 5G shows j2^(CR)ej2^(CR) double mutant plant (left) and inflorescence (right) showingSIM overproliferation and few flowers late in development, respectively.FIG. 5H shows j2^(CR) ej2^(CR) Zi^(CR) triple mutant. Stereoscope images(insets) of j2^(CR) ej2^(CR) Zin^(CR) triple mutants showing massive SIMoverproliferation and no floral termination. FIG. 5I shows j2^(CR) ej2^(CR) Zin^(CR) triple mutant in S. pimpinellifolium as in FIG. 5Hshowing massive SIM overproliferation and no floral termination. Stripedarrowheads indicate successive inflorescences. Scale bars represent 1 cmand 1 mm for photographs and stereoscopic images, respectively.

FIGS. 6A-6D show the exploiting dosage effects of key meristemmaturation genes to improve flower production and fruit yield. FIG. 6Ashows representative inflorescences from different genotypiccombinations of natural and engineered j2 and ej2 mutations in M82. Redarrowheads indicate branching events. FIG. 6B shows the percentage ofinflorescences with 1 to 5 or greater branching events for the indicatedgenotypes. Circled, lower-case letters match genotypes shown in FIG. 6A.Weakly branched genotypes are highlighted with bolded black circles.FIG. 6C shows representative weakly branched inflorescence of as^(classic)/+ heterozygote. FIG. 6D shows the percentage ofinflorescences with branching events for s^(classic)/+,s^(multiflora)/+, and s^(n5568)/+ heterozygous genotypes. Whitearrowheads in FIGS. 6A and 6C mark inflorescence branch points.Nindicates number of inflorescences (FIGS. 6B and 6D). Scale bars inFIGS. 6A and 6C indicate 1 cm.

FIGS. 7A-7K show that s2 inflorescence branching variants are allelic,fail to complement the classical j2 mutant, and are genetically additivewith s. FIGS. 7A-7C show the accessions LA0315 (FIG. 7A), LA3226 (FIG.7B), and the X-ray-induced mutant frondea (FIG. 7C) (Stubbe, 1972)develop highly proliferated inflorescences that bear flowers and fruitswith jointless pedicels (white asterisks). FIGS. 7D-7F show stereoscopeimages of primary meristems in LA0315 (FIG. 7D), LA3226 (FIG. 7E), andfrondea (FIG. 7F), showing the first inflorescence branching event(white arrowhead) at the base of the first flower (F1). SYM: sympodialshoot meristem; L8: leaf 8. FIGS. 7G-7I show representativeinflorescences of F₁ progeny from the crosses LA0315×s2 (FIG. 7G),LA3226×s2 (FIG. 7H), and fro×LA0315 (FIG. 7I) showing all fouraccessions (mutants) are allelic. Scale bars in FIGS. 7A-7C, 7G-7I, and7D-7F indicate 5 cm and 500 μm, respectively. FIG. 7J showsinflorescences of s (left), s2 (middle), and the s s2 higher-ordermutant (right). Greater inflorescence complexity in the s s2higher-order mutant suggests additivity. FIG. 7K shows a complementationtest using an s2-derived jointless mutant plants and the classical j2mutant. Jointed fruits (dotted asterisk) of WT plants and jointlessfruits (white asterisk) of F₁ progeny from a cross of s2-derived j2 andj2 are shown. Scale bar=1 cm.

FIGS. 8A-8C show the rate of meristem maturation in s2 mutants is lessdelayed than in s. FIG. 8A shows the clustering of 2,582 genes that weredynamically expressed during the early (EVM), middle (MVM), and late(LVM) vegetative meristem, the transition meristem (TM) and floralmeristem (FM) stage of meristem maturation in the WT (see STAR Methods).Genes in Cluster 06 and Cluster 08 (solid line boxes) were selected asTM and FM marker genes, respectively. Thick black lines indicate medianexpression with dotted area representing the 5^(th) and 95^(th)quantile. N=number of genes. FIGS. 8B and 8C show WT, s (top), and s2(bottom) z-score normalized expression of TM marker genes in vegetative(VM), transition (TM), and floral (FM) meristem stages. Cluster indotted line boxes and solid line boxes were selected as moderately andstrongly delayed genes, respectively.

FIGS. 9A-9J show that mapping-by-sequencing reveals s2 branching iscaused by mutations in two tomato homologs of the SEPALLATA MADS-boxgenes (J2 and EJ2). FIGS. 9A and 9B show representative images of thephenotypic classes found in the M82×s2 F₂ (FIG. 9A) and S.pimpinellifolium×s2 F₂ populations (FIG. 9B). Asterisks mark jointlesspedicels and arrowheads mark inflorescence branching events. Scalebars=1 cm. FIG. 9C shows segregation ratios of the s2 branchingphenotype in the two F₂ populations. Note that in the M82×s2 F₂, the j2and s2 phenotypes segregated ¼ and 1/16, respectively. FIG. 9D showsmapping-by-sequencing of the loci underlying s2 in an M82×s2 F₂population. Pooled DNA from WT, j2 and s2 plants was sequenced and theratios of the SNP-ratios (s2/M82) between different phenotypic classes(top: s2/WT; middle: s2/j2; bottom: jointless/WT) are shown. FIG. 9Eshows mapping-by-sequencing of the loci underlying s2 in a S.pimpinellifolium×s2 F₂ population. Pooled DNA from WT,j2, and s2 plantswas sequenced and ratios of the SNP-ratios (S.lyc/S.pim) are shown as inFIG. 9D. FIG. 9F shows partial sequence alignment of J2 (Solyc12g038510)from M82, the jointless S. cheesmaniae (S. che) accession LA0166, theclassical j2 accession (LA0315) and the s2 accession (LA4371). A S.cheesmaniae SNP in the second exon leads to a premature stop-codon(asterisk). Allele designated as j2^(stop). From top to bottom,sequences correspond to SEQ ID NOs: 98, 98, 99, 100, and 101. FIG. 9Gshows the CAPS marker PCR genotyping assay for j2^(stop) in accessionsfrom FIG. 9F. Positions of WT and mutant (mut) bands are indicated. FIG.9H shows gene models showing the position of the Copia/Ridertransposable element (TE) insertion in j2^(TE) and the S. cheesmaniaeSNP in j2^(stop). Predicted RNA transcripts are shown below. Thej2^(stop) allele results in a premature stop codon in the second exon.The j2^(TE) allele results in an intronic transcriptional start site andan early stop codon. FIG. 9I shows representative inflorescences of WT,ej2^(w), ej2^(CR), and e j2^(CR)×ej2^(w) F₁ progeny are shown. Scalebar=1 cm. FIG. 9J shows genotyping of s2, LA0315, LA3226,frondea (fro),and WT plants using diagnostic PCR markers for j2^(TE),j2^(stop), andej2^(w). Note that both s2 and LA3226 carry the j2^(TE) and ej2^(w)alleles, whereas LA0315 carries j2^(stop) and ej2^(w). The frondeamutant carries ej2^(TE), however, failed J2 amplification in frondeausing both j2 markers suggest a large structural variant has disruptedthe gene (SV). Band sizes are in kilobase pairs (kbp).

FIGS. 10A-10S show that the three SEP4 genes J2, EJ2 andSolyc04g005320/LIN interact to regulate branching and flowerdevelopment. FIG. 10A shows normalized gene expression (RPKM) for TM5and TM29 (left) and the SEP4 sub-clade (right) in major tissues.

FIG. 10B shows yeast two-hybrid assays showing heteromeric interactionof Solyc04g005320, RIN, J2, and EJ2, and homomeric interaction ofSolyc04g005320, RIN and J2 (3-AT, 3-amino-1,2,4-triazole; L, leucine; T,tryptophan; H, histidine; A, adenine; e.v., empty vector). FIG. 10Cshows the summary of results in FIG. 10B; (−) no interaction; (+)interaction; (++) strong interaction. FIG. 10D shows CRISPR/Cas9targeting of Solyc04g005320. Sequences of Solyc04g005320^(CR) allele 1(a1) and a2 in S. lycopersicum cv. M82 are shown (top). Threeindependent first-generation (T₀) chimeric S. pimpinellifoliumtransgenics were sequenced and 5 reads were obtained per plant (bottom).All sequenced alleles carried mutations, revealing putative biallelic(T₀ #4), homozygous (T₀ #8), and chimeric (T₀ #9) plants. From top tobottom, sequences correspond to SEQ ID NOs: 102-111. FIG. 10E shows thequantification of flowers per inflorescence for WT and 3 independentlin^(CR) T₀ transgenics. N=number of inflorescences. FIG. 10F shows thequantification of internode length between flowers of the same plants asin FIG. 10E. N=number of internodes. FIG. 10G shows representative linmutant plant with elongated and weakly branched inflorescences. Whitearrowheads indicate branch points. Inset shows lin fruit with jointedpedicel. FIG. 10H shows quantification of flowers per inflorescence forWT and lin. N=number of inflorescences. FIG. 10I shows quantification ofinflorescence branching events in WT and lin. FIGS. 10J and 10K showmapping-by-sequencing of the lin mutation in a lin×S. pim F₂ populationto a 0.5 Mbp mapping interval on chromosome 4 containing 80 genesincluding Solyc04g005320. Reads mapping to chromosome 4 indicate atranslocation in Solyc04g005320, which was assayed by PCR (FIG. 10K).The sequence in FIG. 10J corresponds to SEQ ID NO: 112. The WT allele(wt) was amplified with primer-F1 and primer-R2, which bind 5′ and 3′ tothe translocation site, respectively. The lin mutant allele (m) wasamplified with primer-F3, which binds the 3′ border of the translocatedsequence, and primer-R2. FIG. 10L shows semi-quantitative RT-PCR ofSolyc04g005320 in WT and lin showing loss of Solyc04g005320 transcriptin the lin mutant. UBIQUITIN(UBI) was used as control. FIG. 10M showsj2^(CR) lin double mutant with elongated, weakly branched inflorescencesand jointless pedicel (white asterisk). White arrowheads mark branchpoints. FIG. 10N shows ej2^(CR) lin double mutant with longinflorescences, extremely enlarged sepals, and inner floral organdefects (inset). FIG. 10O shows simultaneous targeting of LIN, J2 andE/2 by CRISPR/Cas9 with two single-guide RNAs. sgRNA, Target 1 andTarget 2 on each respective gene model is shown. Note that sgRNA-1targets all three genes. Black arrows indicate forward (F) and reverse(R) primers used for PCR genotyping and sequencing (see STAR Methods).Sequencing results of second-generation (Ti) transgenicj2^(CR) ej2^(CR)lin^(CR) triple mutant plants generated in M82 (top) and S.pimpinellifolium (bottom). All three genes carry homozygous mutations.From top to bottom, sequences correspond to SEQ ID NOs: 113-124. FIG.10P shows CRISPR/Cas9 targeting of LIN in the elite cherry cultivarSweet 100. Sequences of lin^(CR)-allele 1 (a1) and a2 in thefirst-generation (T₀) lin^(CR) plant #1. Five reads were obtained perplant. All sequenced alleles carried mutations, including a complexrearrangement (italicized font). From top to bottom, sequencescorrespond to SEQ ID NOs: 125-127. FIG. 10Q shows representative imagesof Sweet 100 and Sweet 100 lin^(CR) T₀ #1 mutant inflorescences showingdifferent degrees of branching. FIGS. 10R and 10S show quantification offlowers per inflorescence (FIG. 10R) and inflorescence branching events(FIG. 10S) for Sweet 100 and Sweet 100 lin^(CR) T₀#1. N=number ofinflorescences. Bar graphs in FIGS. 10E, 10F., 10H, 10I, 10R, and 10Sshow means (±SEM). P-values determined by two-tailed, two-samplet-tests. Scale bars represent 1 cm.

SEQUENCES

Below is a brief description of certain sequences described herein.

SEQ ID NO: 1 is a nucleic acid sequence of a wild-type Solyc04g005320gene.

SEQ ID NO: 2 is a nucleic acid sequence of a wild-type Solyc04g005320coding sequence.

SEQ ID NO: 3 is a nucleic acid sequence for a mutant Solyc04g005320 geneallele lin^(trans). The border sequences of a translocation site areshown in bold italic letters, with the translocation sequence beingrepresented by the NNNNNN(N*X)NNNNNN sequence.

SEQ ID NO: 4 is a nucleic acid sequence for a mutant Solyc04g005320 geneallele lin^(CR)-allele 1.

SEQ ID NO: 5 is a nucleic acid sequence for a mutant Solyc04g005320 geneallele lin^(CR)-allele 2.

SEQ ID NO: 6 is a nucleic acid sequence of a wild-type Solyc12g038510gene.

SEQ ID NO: 7 is a nucleic acid sequence of a wild-type Solyc12g038510coding sequence.

SEQ ID NO: 8 is a nucleic acid sequence for a mutant Solyc12g038510 geneallele j2^(TE). The border sequences of a transposable element insertionsite are shown in bold italic letters, with the transposable elementsequence being represented by the NNNNNN(N*X)NNNNNN sequence.

SEQ ID NO: 9 is a nucleic acid sequence of a mutant Solyc12g038510 geneallele j2 stop SEQ ID NO: 10 is a nucleic acid sequence for a mutantSolyc04g005320 gene allele j2^(CR)-allele 1.

SEQ ID NO: 11 is a nucleic acid sequence for a mutant Solyc04g005320gene allele j2^(CR)-allele 2.

SEQ ID NO: 12 is a nucleic acid sequence of a wild-type Solyc03g114840gene.

SEQ ID NO: 13 is a nucleic acid sequence of a wild-type Solyc03g114840coding sequence.

SEQ ID NO: 14 is a nucleic acid sequence for a mutant Solyc03g114840gene allele ej2^(W).

SEQ ID NO: 15 is a nucleic acid sequence for a mutant Solyc04g005320gene allele ej2^(CR)-allele 1.

SEQ ID NO: 16 is a nucleic acid sequence for a mutant Solyc04g005320gene allele ej2^(CR)-allele 3.

DETAILED DESCRIPTION

Variation in inflorescence architecture is based on changes in theactivity of meristems, small groups of stem cells located at the tips ofshoots (Kyozuka et al., 2014; Park et al., 2014a). During the transitionto flowering, vegetative meristems gradually mature to a reproductivestate and, depending on the species, terminate immediately in a floweror give rise to a variable number of new inflorescence meristems thatbecome additional flowers or flower-bearing branches (Prusinkiewicz etal., 2007). In domesticated tomato (Solanum lycopersicum) and its wildprogenitor S. pimpinellifolium, a new inflorescence meristem emerges atthe flank of each previous meristem. Several reiterations of thisprocess give rise to inflorescences with multiple flowers arranged in azigzag pattern, resulting in the familiar “tomatoes on the vine”architecture (FIG. 1A)(Park et al., 2012).

Improving tomato inflorescence architecture to boost flower productionand yield has remained surprisingly challenging, despite a rich resourceof wild relatives that develop weakly branched inflorescences with highfertility (Lemmon et al., 2016; Lippman et al., 2008; Park et al., 2012;Zamir, 2001). However, genetic incompatibilities and the challenge oftransferring complex polygenic traits without undesired effects fromlinked genes has precluded exploiting wild species to improveinflorescence architecture (MacArthur and Chiasson, 1947). Anothersource of potentially valuable inflorescence variation is rare naturaland induced highly branched mutants in domesticated germplasm. It waspreviously shown that branching in one of these variants and branchingin a wild species is due to an extended meristem maturation schedule,which allows additional inflorescence meristems to form (Lemmon et al.,2016; Park et al., 2012). This suggested subtle modification of meristemmaturation could provide beneficial changes in inflorescencearchitecture (Park et al., 2014a). Yet, breeders typically selectagainst even moderate branching, primarily due to an imbalance insource-sink relationships that results in high flower abortion and lowfruit production, especially in large-fruited varieties (Stephenson,1981).

In some aspects, the present disclosure relates to the discovery of theidentity of mutations in two closely related MADS-box transcriptionfactor genes, one of which arose during domestication and the otherwithin the last century of crop improvement. Each mutant was selectedseparately based on the phenotype of improved flower morphology andfruit retention traits without knowledge of the locations of themutations and, therefore, the underlying genes affected by themutations. However, combining these two mutants revealed some redundancyin controlling meristem maturation, which caused undesirable branching.Breeders overcame this negative epistasis by selecting suppressors ofbranching, but in so doing limited the potential to improve flowerproduction through weak branching.

As described herein, the identification of the mutations in MADS-boxtranscription factor genes and the dissection of the interaction betweenthe MADS-box genes by Applicants revealed a dosage relationship amongnatural and gene-edited mutations in multiple regulators of meristemmaturation. Combining two or more of the mutations in the MADS-box genesin homozygous and heterozygous combinations allowed for the creation ofa quantitative range of inflorescence types, and the development ofweakly branched hybrids with desirable traits, such as higher flower andfruit production. In particular, data described herein in tomato plantsdemonstrates the utility of mutant MADS-box genes, such as mutant SEP4homologs, and the interaction between such mutant genes to alterinflorescence phenotypes. In particular, mutants of the MADS-box geneSolyc12g038510, mutants of the MADS-box gene Solyc03g114840, and mutantsof the MADS-box gene Solyc04g005320, each of which are homologs ofArabidopsis SEPALLATA4 (SEP4), were shown to be capable of alteringinflorescence phenotypes in tomato plants. Specifically, it was foundthat mixing and matching these mutations in various homozygous andheterozygous combinations resulted in a quantitative range ofinflorescence phenotypes and the development of weakly branched hybridswith higher flower and fruit production.

Accordingly, in some aspects, the present disclosure relates to plants(e.g., Solanaceae plants) comprising one or more mutant MADS-box genessuch as mutant SEPALLATA4 (SEP4) homologs, which may provide a range ofinflorescence phenotypes and may result in improved inflorescencearchitecture and yield.

In some aspects, provided herein are genetically-altered Solanaceaeplants, such as genetically-altered Solanaceae (e.g., Solanumlycopersicum) plants comprising one or more of a mutant Solyc04g005320gene (or a homolog thereof), a mutant Solyc12g038510 gene (or a homologthereof), and a mutant Solyc03g114840 gene (or a homolog thereof), whichexhibit characteristics different from a reference plant such as acorresponding plant that has not been genetically altered (also referredto herein as “wild-type”) or a corresponding plant comprising a nullmutation of one or more of the Solyc04g005320 gene, the Solyc12g038510gene, and the Solyc03g114840 gene. The characteristics include, but arenot limited to, one or more of the following: modified inflorescencearchitecture, modified flower number, higher yield, higher qualityproducts (e.g., fruits), and modified fruit productivity (e.g., modifiedsuch as higher fruit number).

In some embodiments, genetically-altered Solanaceae plants, e.g., tomatoplants (such as Solanum lycopersicum), comprise one or more of a mutantSolyc04g005320 gene (heterozygous or homozygous), a mutantSolyc12g038510 gene (heterozygous or homozygous), and a mutantSolyc03g114840 gene (heterozygous or homozygous). In some embodiments,the plants comprise a variety of combinations of the different mutantalleles, such as, for example, mutant Solyc04g005320 with mutantSolyc12g038510; mutant Solyc04g005320 with mutant Solyc03g114840; ormutant Solyc04g005320 with mutant Solyc12g038510 and mutantSolyc03g114840. The genetically-altered plants may be heterozygotes orhomozygotes and, in some embodiments, may be double heterozygotes,double homozygotes, triple heterozygotes, or triple homozygotes. In someembodiments, such a plant comprises a mutant Solyc04g005320 gene asdescribed herein. In some embodiments, such a plant comprises a mutantSolyc04g005320 gene as described herein and a mutant Solyc12g038510 geneas described herein. In some embodiments, such a plant comprises amutant Solyc04g005320 gene as described herein and a mutantSolyc03g114840 gene as described herein. In some embodiments, such aplant comprises a mutant Solyc04g005320 as described herein with amutant Solyc12g038510 as described herein and a mutant Solyc03g114840 asdescribed herein.

Mutant Solyc04g005320 Gene

Aspects of the disclosure relate to mutants of the Solyc04g005320 gene(or a homolog thereof) as well as plants, plant cells, seeds, andnucleic acids comprising such mutant genes. The Solyc04g005320 gene isalso referred to herein as Long Inflorescence or LIN. The Solyc04g005320gene is a homolog of SEP4 in Arabidopsis.

In some embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc04g005320 gene (or a homolog thereof), such asa hypomorphic allele or null allele, have long inflorescences, e.g.,producing an average of at least 15 flowers (e.g., 9 to 30 flowers) oneach inflorescence per plant. In some embodiments, the number of flowersper inflorescence may vary by variety (e.g. for plum varieties 9-15flowers and for cherry varieties 20-40 flowers). In some embodiments,Solanaceae plants (e.g., Solanum lycopersicum) comprising a mutantSolyc04g005320 gene (or a homolog thereof), such as a hypomorphic alleleor null allele, have longer inflorescences than a plant comprising awild-type Solyc04g005320 gene (or a wild-type homolog thereof). In someembodiments, the mutant Solyc04g005320 gene (or homolog thereof) is ahypomorphic allele that, when crossed to a null allele of theSolyc04g005320 gene (or homolog thereof), does not restore a wild-typeSolyc04g005320 gene (or a wild-type homolog thereof) phenotype (such asproducing an average of 8 flowers (e.g., 6 to 10 flowers) on eachinflorescence per plant). In some embodiments, Solanaceae plants (e.g.,Solanum lycopersicum) comprising a mutant Solyc04g005320 gene (or ahomolog thereof), such as a hypermorphic allele, have shortinflorescences, e.g., producing an average of less than 5 flowers (e.g.,2 to 6 flowers) on each inflorescence per plant. In some embodiments,plants comprising a mutant Solyc04g005320 gene, such as a hypermorphicallele, have shorter inflorescence than a plant comprising a wild-typeSolyc04g005320 gene.

In some embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc04g005320 gene (or a homolog thereof), such asa hypomorphic allele or null allele, have more branches perinflorescence, e.g., producing 2 or more branches per inflorescence. Insome embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc04g005320 gene (or a homolog thereof), such asa hypomorphic allele or null allele, have more branches than a plantcomprising a wild-type Solyc04g005320 gene (or a wild-type homologthereof). In some embodiments, the mutant Solyc04g005320 gene (orhomolog thereof) is a hypomorphic allele that, when crossed to a nullallele of the Solyc04g005320 gene, does not restore a wild-typeSolyc04g005320 gene (or a wild-type homolog thereof) phenotype (such asproducing an average of 1 branch per inflorescence).

In some embodiments, the mutant Solyc04g005320 gene (or homolog thereof)contains a mutation in a regulatory region, a coding region or both(e.g., a missense, nonsense, insertion, deletion, duplication,inversion, indel, or translocation mutation in such a region). In someembodiments, the regulatory region is a promoter. In some embodiments,the mutation in the coding region is in an exon. In some embodiments,the mutation is a translocation in the first intron (e.g., lin^(trans),which contains a translocation in the first intron that eliminatestranscription). In some embodiments, the mutation is a null mutation inwhich the coding sequence has been deleted (e.g., lin^(CR) which is anull allele produced by CRISPR/Cas9).

In some embodiments, the mutant Solyc04g005320 gene (or homolog thereof)is a hypomorphic allele or a null allele. In some embodiments, ahypomorphic allele is an allele that results in an mRNA or proteinexpression level of the gene of interest that is at least 30% lower(e.g., at least 30%, at least 40%, at least 50%, at least 60%, at least70%, at least 80% or at least 90%) than results from an allele of thegene of interest that does not contain the mutation (e.g., a wild-typeallele). As used herein, a “null allele” refers to an allele of a geneof interest in which transcription into RNA does not occur, translationinto a functional protein does not occur or neither occurs due to amutation which may be located within the coding sequence, in aregulatory region of the gene, or in both (e.g., a missense, nonsense,insertion, deletion, duplication, inversion, indel, or translocation).In some embodiments, the null allele is a knock-out allele. As usedherein, a “knock out allele” refers to an allele of a gene in whichtranscription into RNA does not occur, translation into a functionalprotein does not occur or neither occurs as a result of a deletion ofsome portion or all of the coding sequence of the gene, e.g., usinghomologous recombination. One non-limiting approach to creating nullmutations is to use CRISPR-Cas9 mutagenesis to target exons that encodefunctional protein domains or to target a large portion (e.g., at least80%) of the coding sequence (see, e.g., Shi et al. Nature Biotechnology.(2015) 33(6): 661-667 and Online Methods).

In some embodiments, the mutant Solyc04g005320 gene (or homolog thereof)is a hypermorphic allele. In some embodiments, a hypermorphic allele isan allele that results in an mRNA or protein expression level of thegene of interest that is at least 30% greater (e.g., at least 30%, atleast 40%, at least 50%, at least 60%, at least 70%, at least 80%, atleast 90%, at least 100%, at least 200% or more) than results from anallele of the gene of interest that does not contain the mutation (e.g.,a wild-type allele). mRNA and protein levels can be measured using anymethod known in the art or described herein, e.g., using qRT-PCR formRNA levels or an immunoassay for protein levels.

In some embodiments, a Solanaceae plant (e.g., Solanum lycopersicum)comprising the mutant Solyc04g005320 gene, or homolog thereof, (e.g., ahypomorphic, knock-out or null allele described herein) is heterozygousfor the mutant gene. In some embodiments, a Solanaceae plant (e.g.,Solanum lycopersicum) comprising the mutant Solyc04g005320 gene, orhomolog thereof, (e.g., a hypomorphic, knock-out or null alleledescribed herein) is homozygous for the mutant gene.

In some embodiments, the Solyc04g005320 gene homolog (a) has a sequencethat has at least 75%, at least 80%, at least 85%, at least 90%, atleast 95%, at least 96%, at least 97%, at least 98%, or at least 99%identity with the sequence of SEQ ID NO: 1 or 2 and (b) is not a Solanumlycopersicum gene.

In some embodiments, the mutant lin^(trans) gene comprises, for example,a nucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 3; aportion of SEQ ID NO: 3 that exhibits substantially the same activity(e.g., encoding the same polypeptide or substantially the samepolypeptide that has the same activity) as a nucleic acid (e.g., DNA)having the sequence of SEQ ID NO: 3; a nucleic acid (e.g., DNA) havingat least 85%, at least 90%, at least 95%, at least 96%, at least 97%, atleast 98%, or at least 99% identity with the sequence of SEQ ID NO: 3;an orthologue or homologue of the nucleic acid having the sequence ofSEQ ID NO: 3.

In some embodiments, the mutant lin^(CR) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 4 or 5; aportion of SEQ ID NO: 4 or 5 that exhibits substantially the sameactivity (e.g., encoding the same polypeptide or substantially the samepolypeptide that has the same activity) as a nucleic acid (e.g., DNA)having the sequence of SEQ ID NO: 4 or 5; a nucleic acid (e.g., DNA)having at least 85%, at least 90%, at least 95%, at least 96%, at least97%, at least 98%, or at least 99% identity with the sequence of SEQ IDNO: 4 or 5; an orthologue or homologue of the nucleic acid having thesequence of SEQ ID NO: 4 or 5.

Mutant Solyc12g038510 Gene

Other aspects of the disclosure relate to mutants of the Solyc12g038510gene (or a homolog thereof) as well as plants, plant cells, seeds, andnucleic acids comprising such mutant genes. The Solyc12g038510 gene isalso referred to herein as Jointless-2 or J2. The Solyc12g038510 gene isa homolog of SEP4 in Arabidopsis.

In some embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc12g038510 gene (or homolog thereof), such as ahypomorphic allele or null allele, have more branches, e.g., producing 2or more branches per inflorescence. In some embodiments, Solanaceaeplants (e.g., Solanum lycopersicum) comprising a mutant Solyc12g038510gene (or a homolog thereof), such as a hypomorphic allele or nullallele, have more branches than a plant comprising a wild-typeSolyc12g038510 gene. In some embodiments, the mutant Solyc12g038510 gene(or homolog thereof) is a hypomorphic allele that, when crossed to anull allele of the Solyc12g038510 gene (or homolog thereof), does notrestore a wild-type Solyc12g038510 gene (or a wild-type homolog thereof)phenotype (such as producing an average of 1 branch per inflorescence).In some embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc12g038510 gene (or a homolog thereof), such asa hypomorphic allele or null allele, lack the abscission zone on thestems (pedicels) of flowers known as the joint (this absence of theabscission zone is also referred to herein as “jointless pedicels”) orproduce a visible abscission zone (i.e. joint) but abscission does notoccur or requires more force (e.g., hand harvesting) to separate thefruit from the pedicel, providing better fruit retention properties. Insome embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc12g038510 gene (or a homolog thereof), such asa hypomorphic allele or null allele, have more jointless pedicels than aplant comprising a wild-type Solyc12g038510 gene (or a wild-type homologthereof). In some embodiments, the mutant Solyc12g038510 gene (orhomolog thereof) is a hypomorphic allele that, when crossed to a nullallele of the Solyc12g038510 gene (or homolog thereof), does not restorea wild-type Solyc12g038510 gene (or a wild-type homolog thereof)phenotype (such as having a normal abscission zone on the pedicels).

In some embodiments, the mutant Solyc12g038510 gene (or homolog thereof)contains a mutation in a regulatory region, a coding region or both(e.g., a missense, nonsense, insertion, deletion, duplication,inversion, indel, or translocation mutation in such a region). In someembodiments, the regulatory region is a promoter. In some embodiments,the mutation in the coding region is in an exon. In some embodiments,the mutation is in the first intron (e.g., j2^(TE) which contains aCopia/Rider-type transposable element (TE) in the first intron). In someembodiments, the mutation is a nonsense mutation that results in anearly stop codon (e.g., j2^(stop) has an early nonsense mutation). Insome embodiments, the mutation is a null mutation in which the codingsequence has been deleted (e.g., j2^(CR) which is a null allele producedby CRISPR/Cas9).

In some embodiments, the mutant Solyc12g038510 gene (or homolog thereof)is a hypomorphic allele or a null allele. In some embodiments, ahypomorphic allele is an allele that results in an mRNA or proteinexpression level of the gene of interest that is at least 30% lower(e.g., at least 30%, at least 40%, at least 50%, at least 60%, at least70%, at least 80% or at least 90%) than results from an allele of thegene of interest that does not contain the mutation (e.g., a wild-typeallele).

In some embodiments, a Solanaceae plant (e.g., Solanum lycopersicum)comprising the mutant Solyc12g038510 gene, or homolog thereof, (e.g., ahypomorphic, knock-out or null allele described herein) is heterozygousfor the mutant gene. In some embodiments, a Solanaceae plant (e.g.,Solanum lycopersicum) comprising the mutant Solyc12g038510 gene, orhomolog thereof, (e.g., a hypomorphic, knock-out or null alleledescribed herein) is homozygous for the mutant gene.

In some embodiments, the Solyc12g038510 gene homolog (a) has a sequencethat has at least 75%, at least 80%, at least 85%, at least 90%, atleast 95%, at least 96%, at least 97%, at least 98%, or at least 99%identity with the sequence of SEQ ID NO: 6 or 7 and (b) is not a Solanumlycopersicum gene.

In some embodiments, the mutant j2^(TE) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 8; a portionof SEQ ID NO: 8 that exhibits substantially the same activity (e.g.,encoding the same polypeptide or substantially the same polypeptide thathas the same activity) as a nucleic acid (e.g., DNA) having the sequenceof SEQ ID NO: 8; a nucleic acid (e.g., DNA) having at least 85%, atleast 90%, at least 95%, at least 96%, at least 97%, at least 98%, or atleast 99% identity with the sequence of SEQ ID NO: 8; an orthologue orhomologue of the nucleic acid having the sequence of SEQ ID NO: 8.

In some embodiments, the mutant j2^(stop) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 9; a portionof SEQ ID NO: 9 that exhibits substantially the same activity (e.g.,encoding the same polypeptide or substantially the same polypeptide thathas the same activity) as a nucleic acid (e.g., DNA) having the sequenceof SEQ ID NO: 9; a nucleic acid (e.g., DNA) having at least 85%, atleast 90%, at least 95%, at least 96%, at least 97%, at least 98%, or atleast 99% identity with the sequence of SEQ ID NO: 9; an orthologue orhomologue of the nucleic acid having the sequence of SEQ ID NO: 9.

In some embodiments, the mutant j2^(CR) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 10 or 11; aportion of SEQ ID NO: 10 or 11 that exhibits substantially the sameactivity (e.g., encoding the same polypeptide or substantially the samepolypeptide that has the same activity) as a nucleic acid (e.g., DNA)having the sequence of SEQ ID NO: 10 or 11; a nucleic acid (e.g., DNA)having at least 85%, at least 90%, at least 95%, at least 96%, at least97%, at least 98%, or at least 99% identity with the sequence of SEQ IDNO: 10 or 11; an orthologue or homologue of the nucleic acid having thesequence of SEQ ID NO: 10 or 11.

Mutant Solyc03g114840 Gene

Other aspects of the disclosure relate to mutants of the Solyc03g114840gene (or a homolog thereof) as well as plants, plant cells, seeds, andnucleic acids comprising such mutant genes. The Solyc03g114840 gene isalso referred to herein as Enhancer-of-Jointless-2 or EJ2. TheSolyc03g114840 gene is a homolog of SEP4 in Arabidopsis.

In some embodiments, Solanaceae plants (e.g., Solanum lycopersicum)comprising a mutant Solyc03g114840 gene (or a homolog thereof), such asa hypomorphic allele or null allele, have more branches, e.g., producing2 or more branches per inflorescence. In some embodiments, Solanaceaeplants (e.g., Solanum lycopersicum) comprising a mutant Solyc03g114840gene (or a homolog thereof), such as a hypomorphic allele or nullallele, have more branches than a plant comprising a wild-typeSolyc03g114840 gene (or a wild-type homolog thereof). In someembodiments, the mutant Solyc03g114840 gene (or homolog thereof) is ahypomorphic allele that, when crossed to a null allele of theSolyc03g114840 gene (or homolog thereof), does not restore a wild-typeSolyc03g114840 gene (or a wild-type homolog thereof) phenotype (such asproducing an average of 1 branch per inflorescence). In someembodiments, Solanaceae plants (e.g., Solanum lycopersicum) comprising amutant Solyc03g114840 gene (or a homolog thereof), such as a hypomorphicallele or null allele, have long sepals resulting in larger calyxes,e.g., that are an average sepal to petal ratio (sepal length/petallength) of at least 1.2. In some embodiments, Solanaceae plants (e.g.,Solanum lycopersicum) comprising a mutant Solyc03g114840 gene (or ahomolog thereof), such as a hypomorphic allele or null allele, havelonger sepals than a plant comprising a wild-type Solyc03g114840 gene(or a wild-type homolog thereof). In some embodiments, the mutantSolyc03g114840 gene (or homolog thereof) is a hypomorphic allele that,when crossed to a null allele of the Solyc03g114840 gene (or homologthereof), does not restore a wild-type Solyc03g114840 gene (or wild-typehomolog thereof) phenotype (such as having an average sepal to petalratio (sepal length/petal length) of not more than 0.8).

In some embodiments, the mutant Solyc03g114840 gene (or homolog thereof)contains a mutation in a regulatory region, a coding region or both(e.g., a missense, nonsense, insertion, deletion, duplication,inversion, indel, or translocation mutation in such a region). In someembodiments, the regulatory region is a promoter. In some embodiments,the mutation is a null mutation in which the coding sequence has beendeleted (e.g., ej2^(CR) which is a null allele produced by CRISPR/Cas9).In some embodiments, the mutation is an insertion mutation in the 5^(th)intron (e.g., ej2^(W) which is a hypomorphic allele with a 564 bpinsertion in the 5th intron).

In some embodiments, the mutant Solyc03g114840 gene (or homolog thereof)is a hypomorphic allele or a null allele. In some embodiments, ahypomorphic allele is an allele that results in an mRNA or proteinexpression level of the gene of interest that is at least 30% lower(e.g., at least 30%, at least 40%, at least 50%, at least 60%, at least70%, at least 80% or at least 90%) than results from an allele of thegene of interest that does not contain the mutation (e.g., a wild-typeallele).

In some embodiments, a Solanaceae plant (e.g., Solanum lycopersicum)comprising the mutant Solyc03g114840 gene, or homolog thereof, (e.g., ahypomorphic, knock-out or null allele described herein) is heterozygousfor the mutant gene. In some embodiments, a Solanaceae plant (e.g.,Solanum lycopersicum) comprising the mutant Solyc03g114840 gene, orhomolog thereof, (e.g., a hypomorphic, knock-out or null alleledescribed herein) is homozygous for the mutant gene.

In some embodiments, the Solyc03g114840 gene homolog (a) has a sequencethat has at least 75%, at least 80%, at least 85%, at least 90%, atleast 95%, at least 96%, at least 97%, at least 98%, or at least 99%identity with the sequence of SEQ ID NO: 12 or 13 and (b) is not aSolanum lycopersicum gene.

In some embodiments, the mutant ej2^(w) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 14; a portionof SEQ ID NO: 14 that exhibits substantially the same activity (e.g.,encoding the same polypeptide or substantially the same polypeptide thathas the same activity) as a nucleic acid (e.g., DNA) having the sequenceof SEQ ID NO: 14; a nucleic acid (e.g., DNA) having at least 85%, atleast 90%, at least 95%, at least 96%, at least 97%, at least 98%, or atleast 99% identity with the sequence of SEQ ID NO: 14; an orthologue orhomologue of the nucleic acid having the sequence of SEQ ID NO: 14.

In some embodiments, the mutant ej2^(CR) gene comprises, for example, anucleic acid (e.g., DNA) having the sequence of SEQ ID NO: 15 or 16; aportion of SEQ ID NO: 15 or 16 that exhibits substantially the sameactivity (e.g., encoding the same polypeptide or substantially the samepolypeptide that has the same activity) as a nucleic acid (e.g., DNA)having the sequence of SEQ ID NO: 15 or 16; a nucleic acid (e.g., DNA)having at least 85%, at least 90%, at least 95%, at least 96%, at least97%, at least 98%, or at least 99% identity with the sequence of SEQ IDNO: 15 or 16; an orthologue or homologue of the nucleic acid having thesequence of SEQ ID NO: 15 or 16.

Solanaceae Plants Comprising Mutant Genes

Higher yield, higher quality products (e.g., fruits) and products (e.g.,fruits) with different compositions (e.g., brix, also known as enhancedsoluble solids or sugar concentration in the fruits), can be manipulatedin a wide variety of types of Solanaceae plants that comprise a mutantgene, such as a mutant Solyc04g005320 gene (or homolog thereof), amutant Solyc12g038510 gene (or homolog thereof), or a mutantSolyc03g114840 gene (or homolog thereof); or two mutant genes, such asboth a mutant Solyc04g005320 gene (or homolog thereof) and a mutantSolyc12g038510 gene (or homolog thereof), both a mutant Solyc04g005320gene (or homolog thereof) and a mutant Solyc03g114840 gene (or homologthereof), or both a mutant Solyc12g038510 gene (or homolog thereof) anda mutant Solyc03g114840 gene (or homolog thereof); or three mutantgenes, such as a mutant Solyc04g005320 gene (or homolog thereof), amutant Solyc12g038510 gene (or homolog thereof), and a mutantSolyc03g114840 gene (or homolog thereof). In some embodiments, theSolanaceae plant is a genetically-altered Solanaceae plant. In someembodiments, a “genetically-altered” plant includes a plant that has hadintroduced into it (or introduced into a plant that is used to producethe plant, such as introduced into a parental line) at least onemutation by chemical or physical means (e.g., using CRISPR/Cas9,chemical mutagenesis, radiation, Agrobacterium-mediated recombination,viral-vector mediated recombination, or transposon mutagenesis).

The mutant Solyc04g005320 gene (or homolog thereof) can be any of themutant Solyc04g005320 genes (or homologs thereof) described herein. Themutant Solyc12g038510 gene (or homolog thereof) can be any of the mutantSolyc12g038510 genes (or homologs thereof) described herein. The mutantSolyc03g114840 gene (or homolog thereof) can be any of the mutantSolyc03g114840 genes (or homologs thereof) described herein.

The genetically-altered Solanaceae plant can be, for example, inbred,isogenic or hybrid, as long as the plant comprises a mutant gene, suchas a mutant Solyc04g005320 gene (or homolog thereof), a mutantSolyc12g038510 gene (or homolog thereof), or a mutant Solyc03g114840gene (or homolog thereof); or two mutant genes, such as both a mutantSolyc04g005320 gene (or homolog thereof) and a mutant Solyc12g038510gene (or homolog thereof), both a mutant Solyc04g005320 gene (or homologthereof) and a mutant Solyc03g114840 gene (or homolog thereof), or botha mutant Solyc12g038510 gene (or homolog thereof) and a mutantSolyc03g114840 gene (or homolog thereof); or three mutant genes, such asa mutant Solyc04g005320 gene (or homolog thereof), a mutantSolyc12g038510 gene (or homolog thereof), and a mutant Solyc03g114840gene (or homolog thereof).

Plants in the Solanaceae family include, e.g., tomato, potato, eggplant,petunia, tobacco, and pepper. In some embodiments, the Solanaceae plantis a tomato plant. In some embodiments, the Solanaceae plant, e.g.tomato plant, is not a variety.

In some embodiments, the genetically-altered Solanaceae plant comprisesone wild-type (WT) copy of the SOLYC04G005320 gene (or homolog thereof)and one mutant copy of the Solyc04g005320 gene (or homolog thereof) asdescribed herein (is heterozygous for the mutant Solyc04g005320 gene orhomolog thereof). In some embodiments, the Solanaceae plant comprisestwo copies of a mutant Solyc04g005320 gene (or homolog thereof) asdescribed herein (is homozygous for the mutant Solyc04g005320 gene orhomolog thereof). In some embodiments, the Solanaceae plant comprises afirst mutant Solyc04g005320 gene (or homolog thereof) as describedherein and a second mutant Solyc04g005320 gene (or homolog thereof) asdescribed herein, wherein the first mutant Solyc04g005320 gene (orhomolog thereof) and the second mutant Solyc04g005320 gene (or homologthereof) are different. In some embodiments, the Solanaceae plantcomprises one copy of a mutant Solyc04g005320 gene (or homolog thereof)as described herein and one copy of a mutant Solyc12g038510 gene (orhomolog thereof) as described herein (is heterozygous for the mutantSolyc04g005320 gene, or homolog thereof, and heterozygous for the mutantSolyc12g038510 gene, or homolog thereof). In some embodiments, theSolanaceae plant comprises one copy of a mutant Solyc04g005320 gene (orhomolog thereof) as described herein and two copies of a mutantSolyc12g038510 gene (or homolog thereof) as described herein (isheterozygous for the mutant Solyc04g005320 gene, or homolog thereof andhomozygous for the mutant Solyc12g038510 gene, or homolog thereof). Insome embodiments, the Solanaceae plant comprises two copies of a mutantSolyc04g005320 gene (or homolog thereof) as described herein and twocopies of a mutant Solyc12g038510 gene (or homolog thereof) as describedherein (is homozygous for the mutant Solyc04g005320 gene, or homologthereof, and homozygous for the mutant Solyc12g038510 gene, or homologthereof).

In some embodiments, the genetically-altered Solanaceae plant comprisesone WT copy of a SOLYC03G114840 gene (or homolog thereof) and one mutantcopy of a Solyc03g114840 gene (or homolog thereof) as described herein(is heterozygous for the mutant Solyc03g114840 gene, or homologthereof). In some embodiments, the Solanaceae plant comprises two copiesof a mutant Solyc03g114840 gene (or homolog thereof) as described herein(is homozygous for the mutant Solyc03g114840 gene or homolog thereof).In some embodiments, the Solanaceae plant comprises one copy of a mutantSolyc03g114840 gene (or homolog thereof) as described herein and onecopy of a mutant Solyc04g005320 gene (or homolog thereof) as describedherein (is heterozygous for the mutant Solyc03g114840 gene, or homologthereof, and heterozygous for the mutant Solyc04g005320 gene, or homologthereof). In some embodiments, the Solanaceae plant comprises one copyof a mutant Solyc03g114840 gene (or homolog thereof) as described hereinand two copies of a mutant Solyc04g005320 gene (or homolog thereof) asdescribed herein (is heterozygous for the mutant Solyc03g114840 gene, orhomolog thereof, and homozygous for the mutant Solyc04g005320 gene, orhomolog thereof). In some embodiments, the Solanaceae plant comprisestwo copies of a mutant Solyc03g114840 gene (or homolog thereof) asdescribed herein and two copies of a mutant Solyc04g005320 gene (orhomolog thereof) as described herein (is homozygous for the mutantSolyc03g114840 gene, or homolog thereof, and homozygous for the mutantSolyc04g005320 gene, or homolog thereof).

In some embodiments, the genetically-altered Solanaceae plant comprisesone WT copy of a SOLYC03G114840 gene and one mutant copy of aSolyc03g114840 gene as described herein (is heterozygous for the mutantSolyc03g114840 gene) and comprises one WT copy of the SOLYC12G038510gene and one mutant copy of the Solyc12g038510 gene as described herein(is heterozygous for the mutant Solyc12g038510 gene). In someembodiments, the Solanaceae plant comprises two copies of a mutantSolyc03g114840 gene as described herein (is homozygous for the mutantSolyc03g114840 gene) and comprises two copies of a mutant Solyc12g038510gene as described herein (is homozygous for the mutant Solyc12g038510gene). In some embodiments, the Solanaceae plant comprising a mutantSolyc03g114840 gene (one or two copies) as described herein and a mutantSolyc12g038510 gene (one or two copies) further comprises one copy of amutant Solyc04g005320 gene as described herein (is heterozygous orhomozygous for the mutant Solyc03g114840 gene and the mutantSolyc12g038510 gene and heterozygous for the mutant Solyc04g005320gene). In some embodiments, the Solanaceae plant further comprises twocopies of a mutant Solyc04g005320 gene as described herein (ishomozygous for the mutant Solyc04g005320 gene).

Other, non-limiting example genotype combinations which a Solanaceae(e.g., Solanum lycopersicum) plant may comprise are displayed inTable 1. The combinations in Table 1 may also be with homologs of thegenes.

TABLE 1 Example genotype combinations. Combination Solyc12g038510Solyc03g114840 Solyc04g005320 No. (J2) Genotype (EJ2) Genotype (LIN)Genotype 1 j2^(TE)/j2^(TE) ej2^(W)/ej2^(W) lin^(trans)/lin^(trans) 2j2^(TE)/j2^(TE) ej2^(W)/+   lin^(trans)/lin^(trans) 3 j2^(TE)/j2^(TE)+/+ lin^(trans)/lin^(trans) 4 j2^(TE)/j2^(TE) ej2^(W)/ej2^(W)lin^(trans)/+    5 j2^(TE)/j2^(TE) ej2^(W)/+   lin^(trans)/+    6j2^(TE)/j2^(TE) +/+ lin^(trans)/+    7 j2^(TE)/j2^(TE) ej2^(W)/ej2^(W)+/+ 8 j2^(TE)/j2^(TE) ej2^(W)/+   +/+ 9 j2^(TE)/j2^(TE) +/+ +/+ 10j2^(stop)/j2^(stop) ej2^(W)/ej2^(W) lin^(trans)/lin^(trans) 11j2^(stop)/j2^(stop) ej2^(W)/+   lin^(trans)/lin^(trans) 12j2^(stop)/j2^(stop) +/+ lin^(trans)/lin^(trans) 13 j2^(stop)/j2^(stop)ej2^(W)/ej2^(W) lin^(trans)/+    14 j2^(stop)/j2^(stop) ej2^(W)/+  lin^(trans)/+    15 j2^(stop)/j2^(stop) +/+ lin^(trans)/+    16j2^(stop)/j2^(stop) ej2^(W)/ej2^(W) +/+ 17 j2^(stop)/j2^(stop)ej2^(W)/+   +/+ 18 j2^(stop)/j2^(stop) +/+ +/+ 19 j2^(CR)/j2^(CR)ej2^(W)/ej2^(W) lin^(trans)/lin^(trans) 20 j2^(CR)/j2^(CR) ej2^(W)/+  lin^(trans)/lin^(trans) 21 j2^(CR)/j2^(CR) +/+ lin^(trans)/lin^(trans)22 j2^(CR)/j2^(CR) ej2^(W)/ej2^(W) lin^(trans)/+    23 j2^(CR)/j2^(CR)ej2^(W)/+   lin^(trans)/+    24 j2^(CR)/j2^(CR) +/+ lin^(trans)/+    25j2^(CR)/j2^(CR) ej2^(W)/ej2^(W) +/+ 26 j2^(CR)/j2^(CR) ej2^(W)/+   +/+27 j2^(CR)/j2^(CR) +/+ +/+ 28 j2^(TE)/+   ej2^(W)/ej2^(W)lin^(trans)/lin^(trans) 29 j2^(TE)/+   ej2^(W)/+  lin^(trans)/lin^(trans) 30 j2^(TE)/+   +/+ lin^(trans)/lin^(trans) 31j2^(TE)/+   ej2^(W)/ej2^(W) lin^(trans)/+    32 j2^(TE)/+   ej2^(W)/+  lin^(trans)/+    33 j2^(TE)/+   +/+ lin^(trans)/+    34 j2^(TE)/+  ej2^(W)/ej2^(W) +/+ 35 j2^(TE)/+   ej2^(W)/+   +/+ 36 j2^(TE)/+   +/++/+ 37 j2^(stop)/+    ej2^(W)/ej2^(W) lin^(trans)/lin^(trans) 38j2^(stop)/+    ej2^(W)/+   lin^(trans)/lin^(trans) 39 j2^(stop)/+    +/+lin^(trans)/lin^(trans) 40 j2^(stop)/+    ej2^(W)/ej2^(W)lin^(trans)/+    41 j2^(stop)/+    ej2^(W)/+   lin^(trans)/+    42j2^(stop)/+    +/+ lin^(trans)/+    43 j2^(stop)/+    ej2^(W)/ej2^(W)+/+ 44 j2^(stop)/+    ej2^(W)/+   +/+ 45 j2^(stop)/+    +/+ +/+ 46j2^(CR)/+   ej2^(W)/ej2^(W) lin^(trans)/lin^(trans) 47 j2^(CR)/+  ej2^(W)/+   lin^(trans)/lin^(trans) 48 j2^(CR)/+   +/+lin^(trans)/lin^(trans) 49 j2^(CR)/+   ej2^(W)/ej2^(W) lin^(trans)/+   50 j2^(CR)/+   ej2^(W)/+   lin^(trans)/+    51 j2^(CR)/+   +/+lin^(trans)/+    52 j2^(CR)/+   ej2^(W)/ej2^(W) +/+ 53 j2^(CR)/+  ej2^(W)/+   +/+ 54 j2^(CR)/+   +/+ +/+ 55 +/+ ej2^(W)/ej2^(W)lin^(trans)/lin^(trans) 56 +/+ ej2^(W)/+   lin^(trans)/lin^(trans) 57+/+ +/+ lin^(trans)/lin^(trans) 58 +/+ ej2^(W)/ej2^(W) lin^(trans)/+   59 +/+ ej2^(W)/+   lin^(trans)/+    60 +/+ +/+ lin^(trans)/+    61 +/+ej2^(W)/ej2^(W) +/+ 62 +/+ ej2^(W)/+   +/+ 63 +/+ +/+ +/+ 64j2^(TE)/j2^(TE) ej2^(CR)/ej2^(CR) lin^(trans)/lin^(trans) 65j2^(TE)/j2^(TE) ej2^(CR)/+     lin^(trans)/lin^(trans) 66j2^(TE)/j2^(TE) +/+ lin^(trans)/lin^(trans) 67 j2^(TE)/j2^(TE)ej2^(CR)/ej2^(CR) lin^(trans)/+    68 j2^(TE)/j2^(TE) ej2^(CR)/+    lin^(trans)/+    69 j2^(TE)/j2^(TE) +/+ lin^(trans)/+    70j2^(TE)/j2^(TE) ej2^(CR)/ej2^(CR) +/+ 71 j2^(TE)/j2^(TE) ej2^(CR)/+    +/+ 72 j2^(TE)/j2^(TE) +/+ +/+ 73 j2^(stop)/j2^(stop) ej2^(CR)/ej2^(CR)lin^(trans)/lin^(trans) 74 j2^(stop)/j2^(stop) ej2^(CR)/+    lin^(trans)/lin^(trans) 75 j2^(stop)/j2^(stop) +/+lin^(trans)/lin^(trans) 76 j2^(stop)/j2^(stop) ej2^(CR)/ej2^(CR)lin^(trans)/+    77 j2^(stop)/j2^(stop) ej2^(CR)/+     lin^(trans)/+   78 j2^(stop)/j2^(stop) +/+ lin^(trans)/+    79 j2^(stop)/j2^(stop)ej2^(CR)/ej2^(CR) +/+ 80 j2^(stop)/j2^(stop) ej2^(CR)/+     +/+ 81j2^(stop)/j2^(stop) +/+ +/+ 82 j2^(CR)/j2^(CR) ej2^(CR)/ej2^(CR)lin^(trans)/lin^(trans) 83 j2^(CR)/j2^(CR) ej2^(CR)/+    lin^(trans)/lin^(trans) 84 j2^(CR)/j2^(CR) +/+ lin^(trans)/lin^(trans)85 j2^(CR)/j2^(CR) ej2^(CR)/ej2^(CR) lin^(trans)/+    86 j2^(CR)/j2^(CR)ej2^(CR)/+     lin^(trans)/+    87 j2^(CR)/j2^(CR) +/+ lin^(trans)/+   88 j2^(CR)/j2^(CR) ej2^(CR)/ej2^(CR) +/+ 89 j2^(CR)/j2^(CR)ej2^(CR)/+     +/+ 90 j2^(CR)/j2^(CR) +/+ +/+ 91 j2^(TE)/+  ej2^(CR)/ej2^(CR) lin^(trans)/lin^(trans) 92 j2^(TE)/+   ej2^(CR)/+    lin^(trans)/lin^(trans) 93 j2^(TE)/+   +/+ lin^(trans)/lin^(trans) 94j2^(TE)/+   ej2^(CR)/ej2^(CR) lin^(trans)/+    95 j2^(TE)/+  ej2^(CR)/+     lin^(trans)/+    96 j2^(TE)/+   +/+ lin^(trans)/+    97j2^(TE)/+   ej2^(CR)/ej2^(CR) +/+ 98 j2^(TE)/+   ej2^(CR)/+     +/+ 99j2^(TE)/+   +/+ +/+ 100 j2^(stop)/+    ej2^(CR)/ej2^(CR)lin^(trans)/lin^(trans) 101 j2^(stop)/+    ej2^(CR)/+    lin^(trans)/lin^(trans) 102 j2^(stop)/+    +/+ lin^(trans)/lin^(trans)103 j2^(stop)/+    ej2^(CR)/ej2^(CR) lin^(trans)/+    104 j2^(stop)/+   ej2^(CR)/+     lin^(trans)/+    105 j2^(stop)/+    +/+ lin^(trans)/+   106 j2^(stop)/+    ej2^(CR)/ej2^(CR) +/+ 107 j2^(stop)/+   ej2^(CR)/+     +/+ 108 j2^(stop)/+    +/+ +/+ 109 j2^(CR)/+  ej2^(CR)/ej2^(CR) lin^(trans)/lin^(trans) 110 j2^(CR)/+   ej2^(CR)/+    lin^(trans)/lin^(trans) 111 j2^(CR)/+   +/+ lin^(trans)/lin^(trans) 112j2^(CR)/+   ej2^(CR)/ej2^(CR) lin^(trans)/+    113 j2^(CR)/+  ej2^(CR)/+     lin^(trans)/+    114 j2^(CR)/+   +/+ lin^(trans)/+    115j2^(CR)/+   ej2^(CR)/ej2^(CR) +/+ 116 j2^(CR)/+   ej2^(CR)/+     +/+ 117j2^(CR)/+   +/+ +/+ 118 +/+ ej2^(CR)/ej2^(CR) lin^(trans)/lin^(trans)119 +/+ ej2^(CR)/+     lin^(trans)/lin^(trans) 120 +/+ +/+lin^(trans)/lin^(trans) 121 +/+ ej2^(CR)/ej2^(CR) lin^(trans)/+    122+/+ ej2^(CR)/+     lin^(trans)/+    123 +/+ +/+ lin^(trans)/+    124 +/+ej2^(CR)/ej2^(CR) +/+ 125 +/+ ej2^(CR)/+     +/+ 126 j2^(TE)/j2^(TE)ej2^(W)/ej2^(W) lin^(CR)/lin^(CR) 127 j2^(TE)/j2^(TE) ej2^(W)/+  lin^(CR)/lin^(CR) 128 j2^(TE)/j2^(TE) +/+ lin^(CR)/lin^(CR) 129j2^(TE)/j2^(TE) ej2^(W)/ej2^(W) lin^(CR)/+    130 j2^(TE)/j2^(TE)ej2^(W)/+   lin^(CR)/+    131 j2^(TE)/j2^(TE) +/+ lin^(CR)/+    132j2^(TE)/j2^(TE) ej2^(W)/ej2^(W) +/+ 133 j2^(TE)/j2^(TE) ej2^(W)/+   +/+134 j2^(TE)/j2^(TE) +/+ +/+ 135 j2^(stop)/j2^(stop) ej2^(W)/ej2^(W)lin^(CR)/lin^(CR) 136 j2^(stop)/j2^(stop) ej2^(W)/+   lin^(CR)/lin^(CR)137 j2^(stop)/j2^(stop) +/+ lin^(CR)/lin^(CR) 138 j2^(stop)/j2^(stop)ej2^(W)/ej2^(W) lin^(CR)/+    139 j2^(stop)/j2^(stop) ej2^(W)/+  lin^(CR)/+    140 j2^(stop)/j2^(stop) +/+ lin^(CR)/+    141j2^(stop)/j2^(stop) ej2^(W)/ej2^(W) +/+ 142 j2^(stop)/j2^(stop)ej2^(W)/+   +/+ 143 j2^(stop)/j2^(stop) +/+ +/+ 144 j2^(CR)/j2^(CR)ej2^(W)/ej2^(W) lin^(CR)/lin^(CR) 145 j2^(CR)/j2^(CR) ej2^(W)/+  lin^(CR)/lin^(CR) 146 j2^(CR)/j2^(CR) +/+ lin^(CR)/lin^(CR) 147j2^(CR)/j2^(CR) ej2^(W)/ej2^(W) lin^(CR)/+    148 j2^(CR)/j2^(CR)ej2^(W)/+   lin^(CR)/+    149 j2^(CR)/j2^(CR) +/+ lin^(CR)/+    150j2^(CR)/j2^(CR) ej2^(W)/ej2^(W) +/+ 151 j2^(CR)/j2^(CR) ej2^(W)/+   +/+152 j2^(CR)/j2^(CR) +/+ +/+ 153 j2^(TE)/+   ej2^(W)/ej2^(W)lin^(CR)/lin^(CR) 154 j2^(TE)/+   ej2^(W)/+   lin^(CR)/lin^(CR) 155j2^(TE)/+   +/+ lin^(CR)/lin^(CR) 156 j2^(TE)/+   ej2^(W)/ej2^(W)lin^(CR)/+    157 j2^(TE)/+   ej2^(W)/+   lin^(CR)/+    158 j2^(TE)/+  +/+ lin^(CR)/+    159 j2^(TE)/+   ej2^(W)/ej2^(W) +/+ 160 j2^(TE)/+  ej2^(W)/+   +/+ 161 j2^(TE)/+   +/+ +/+ 162 j2^(stop)/+   ej2^(W)/ej2^(W) lin^(CR)/lin^(CR) 163 j2^(stop)/+    ej2^(W)/+  lin^(CR)/lin^(CR) 164 j2^(stop)/+    +/+ lin^(CR)/lin^(CR) 165j2^(stop)/+    ej2^(W)/ej2^(W) lin^(CR)/+    166 j2^(stop)/+   ej2^(W)/+   lin^(CR)/+    167 j2^(stop)/+    +/+ lin^(CR)/+    168j2^(stop)/+    ej2^(W)/ej2^(W) +/+ 169 j2^(stop)/+    ej2^(W)/+   +/+170 j2^(stop)/+    +/+ +/+ 171 j2^(CR)/+   ej2^(W)/ej2^(W)lin^(CR)/lin^(CR) 172 j2^(CR)/+   ej2^(W)/+   lin^(CR)/lin^(CR) 173j2^(CR)/+   +/+ lin^(CR)/lin^(CR) 174 j2^(CR)/+   ej2^(W)/ej2^(W)lin^(CR)/+    175 j2^(CR)/+   ej2^(W)/+   lin^(CR)/+    176 j2^(CR)/+  +/+ lin^(CR)/+    177 j2^(CR)/+   ej2^(W)/ej2^(W) +/+ 178 j2^(CR)/+  ej2^(W)/+   +/+ 179 j2^(CR)/+   +/+ +/+ 180 +/+ ej2^(W)/ej2^(W)lin^(CR)/lin^(CR) 181 +/+ ej2^(W)/+   lin^(CR)/lin^(CR) 182 +/+ +/+lin^(CR)/lin^(CR) 183 +/+ ej2^(W)/ej2^(W) lin^(CR)/+    184 +/+ej2^(W)/+   lin^(CR)/+    185 +/+ +/+ lin^(CR)/+    186 +/+ej2^(W)/ej2^(W) +/+ 187 +/+ ej2^(W)/+   +/+ 188 +/+ +/+ +/+ 189j2^(TE)/j2^(TE) ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 190 j2^(TE)/j2^(TE)ej2^(CR)/+     lin^(CR)/lin^(CR) 191 j2^(TE)/j2^(TE) +/+lin^(CR)/lin^(CR) 192 j2^(TE)/j2^(TE) ej2^(CR)/ej2^(CR) lin^(CR)/+   193 j2^(TE)/j2^(TE) ej2^(CR)/+     lin^(CR)/+    194 j2^(TE)/j2^(TE) +/+lin^(CR)/+    195 j2^(TE)/j2^(TE) ej2^(CR)/ej2^(CR) +/+ 196j2^(TE)/j2^(TE) ej2^(CR)/+     +/+ 197 j2^(TE)/j2^(TE) +/+ +/+ 198j2^(stop)/j2^(stop) ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 199j2^(stop)/j2^(stop) ej2^(CR)/+     lin^(CR)/lin^(CR) 200j2^(stop)/j2^(stop) +/+ lin^(CR)/lin^(CR) 201 j2^(stop)/j2^(stop)ej2^(CR)/ej2^(CR) lin^(CR)/+    202 j2^(stop)/j2^(stop) ej2^(CR)/+    lin^(CR)/+    203 j2^(stop)/j2^(stop) +/+ lin^(CR)/+    204j2^(stop)/j2^(stop) ej2^(CR)/ej2^(CR) +/+ 205 j2^(stop)/j2^(stop)ej2^(CR)/+     +/+ 206 j2^(stop)/j2^(stop) +/+ +/+ 207 j2^(CR)/j2^(CR)ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 208 j2^(CR)/j2^(CR) ej2^(CR)/+    lin^(CR)/lin^(CR) 209 j2^(CR)/j2^(CR) +/+ lin^(CR)/lin^(CR) 210j2^(CR)/j2^(CR) ej2^(CR)/ej2^(CR) lin^(CR)/+    211 j2^(CR)/j2^(CR)ej2^(CR)/+     lin^(CR)/+    212 j2^(CR)/j2^(CR) +/+ lin^(CR)/+    213j2^(CR)/j2^(CR) ej2^(CR)/ej2^(CR) +/+ 214 j2^(CR)/j2^(CR) ej2^(CR)/+    +/+ 215 j2^(CR)/j2^(CR) +/+ +/+ 216 j2^(TE)/+   ej2^(CR)/ej2^(CR)lin^(CR)/lin^(CR) 217 j2^(TE)/+   ej2^(CR)/+     lin^(CR)/lin^(CR) 218j2^(TE)/+   +/+ lin^(CR)/lin^(CR) 219 j2^(TE)/+   ej2^(CR)/ej2^(CR)lin^(CR)/+    220 j2^(TE)/+   ej2^(CR)/+     lin^(CR)/+    221j2^(TE)/+   +/+ lin^(CR)/+    222 j2^(TE)/+   ej2^(CR)/ej2^(CR) +/+ 223j2^(TE)/+   ej2^(CR)/+     +/+ 224 j2^(TE)/+   +/+ +/+ 225j2^(stop)/+    ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 226 j2^(stop)/+   ej2^(CR)/+     lin^(CR)/lin^(CR) 227 j2^(stop)/+    +/+lin^(CR)/lin^(CR) 228 j2^(stop)/+    ej2^(CR)/ej2^(CR) lin^(CR)/+    229j2^(stop)/+    ej2^(CR)/+     lin^(CR)/+    230 j2^(stop)/+    +/+lin^(CR)/+    231 j2^(stop)/+    ej2^(CR)/ej2^(CR) +/+ 232j2^(stop)/+    ej2^(CR)/+     +/+ 233 j2^(stop)/+    +/+ +/+ 234j2^(CR)/+   ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 235 j2^(CR)/+  ej2^(CR)/+     lin^(CR)/lin^(CR) 236 j2^(CR)/+   +/+ lin^(CR)/lin^(CR)237 j2^(CR)/+   ej2^(CR)/ej2^(CR) lin^(CR)/+    238 j2^(CR)/+  ej2^(CR)/+     lin^(CR)/+    239 j2^(CR)/+   +/+ lin^(CR)/+    240j2^(CR)/+   ej2^(CR)/ej2^(CR) +/+ 241 j2^(CR)/+   ej2^(CR)/+     +/+ 242j2^(CR)/+   +/+ +/+ 243 +/+ ej2^(CR)/ej2^(CR) lin^(CR)/lin^(CR) 244 +/+ej2^(CR)/+     lin^(CR)/lin^(CR) 245 +/+ +/+ lin^(CR)/lin^(CR) 246 +/+ej2^(CR)/ej2^(CR) lin^(CR)/+    247 +/+ ej2^(CR)/+     lin^(CR)/+    248+/+ +/+ lin^(CR)/+    249 +/+ ej2^(CR)/ej2^(CR) +/+ 250 +/+ej2^(CR)/+     +/+

Solanaceae plant cells are also contemplated herein. A Solanaceae plantcell may comprise any genotype described herein, e.g., as shown withoutlimitation in Table 1, in the context of the Solanaceae plant (e.g., aSolanaceae plant cell heterozygous for a mutant Solyc03g114840 gene, ora homolog thereof, and a mutant Solyc12g038510 gene, or a homologthereof, or a Solanaceae plant cell homozygous for a mutantSolyc12g038510 gene, or a homolog thereof, and a mutant Solyc04g005320gene, or a homolog thereof). In some embodiments, the Solanaceae plantcell is isolated. In some embodiments, the Solanaceae plant cell is anon-replicating plant cell.

In some embodiments, any of the Solanaceae plants described herein mayan altered phenotype compared to a WT Solanaceae plant (e.g., aSolanaceae plant comprising two copies or one copy of the correspondingWT gene). In some embodiments, any of the Solanaceae plants describedherein have a higher yield than a corresponding WT Solanaceae plant. Insome embodiments, any of the Solanaceae plants described herein have oneor more of the following characteristics: longer sepals, larger calyxes,a different fruit shape, fewer branches, jointless pedicels, longinflorescences, or larger fruit compared to a corresponding WTSolanaceae plant. In some embodiments, such characteristics areappealing to consumers (e.g., products of the Solanaceae plant lookfresher) and are advantageous for growers (e.g., products of theSolanaceae plant stay attached to the plant for a longer period oftime).

Food products are also contemplated herein. Such food products comprisea Solanaceae plant part, such as a fruit (e.g., a tomato fruit).Non-limiting examples of food products include sauces (e.g., tomatosauce or ketchup), purees, pastes, juices, canned fruits, and soups.Food products may be produced or producible by using methods known inthe art.

Isolated polynucleotides are also described herein, including WT andmutant alleles of the Solyc04g005320 gene, or a homolog thereof, e.g.,lin^(trans) and lin^(CR). Isolated polynucleotides including WT andmutant alleles of the Solyc12g038510 gene, or a homolog thereof, arealso contemplated, e.g., j2^(CR) j2^(TE) and j2^(stop). Isolatedpolynucleotides including WT and mutant alleles of the Solyc03g114840gene, or a homolog thereof, are also contemplated, e.g., ej2^(CR) andej2^(W).

Isolated polynucleotides can comprise, for example, a nucleic acid(e.g., DNA) having the sequence of SEQ ID NO: 3, 4, 5, 8, 9, 10, 11, 14,15 or 16; a portion of SEQ ID NO: 3, 4, 5, 8, 9, 10, 11, 14, 15 or 16that exhibits substantially the same activity as a nucleic acid (e.g.,DNA) having the sequence of SEQ ID NO: 3, 4, 5, 8, 9, 10, 11, 14, 15 or16; a nucleic acid (e.g., DNA) having at least 85%, at least 90%, atleast 95%, at least 96%, at least 97%, at least 98%, or at least 99%identity with the sequence of SEQ ID NO: 3, 4, 5, 8, 9, 10, 11, 14, 15or 16; an orthologue or homologue of the nucleic acid having thesequence of SEQ ID NO: 3, 4, 5, 8, 9, 10, 11, 14, 15 or 16. In someembodiments, the isolated polynucleotide is a cDNA. Such isolatedpolynucleotides can be used, for example, in methods of producinggenetically-altered plants.

Other aspects of the disclosure relate to seeds for producing aSolanaceae plant as described herein, e.g., a mutant Solyc04g005320 gene(or a homolog thereof), a mutant Solyc12g038510 gene (or a homologthereof), or a mutant Solyc03g114840 gene (or a homolog thereof).

Methods of Producing Plants

In other aspects, the disclosure provides methods for producing agenetically-altered Solanaceae plant. In some embodiments, the methodcomprises introducing a mutation into a Solyc04g005320 gene (or ahomolog thereof), into a Solyc12g038510 gene (or a homolog thereof), orinto a Solyc03g114840 gene (or a homolog thereof) in the Solanaceaeplant, thereby producing a genetically-altered Solanaceae plantcontaining a mutant version of the gene. In some embodiments, the methodcomprises introducing a mutation into a Solyc04g005320 gene (or ahomolog thereof), into a Solyc12g038510 gene (or a homolog thereof), orinto a Solyc03g114840 gene (or a homolog thereof) in the Solanaceaeplant part, maintaining the plant part under conditions and forsufficient time for production of a genetically-altered Solanaceaeplant, thereby producing a genetically-altered Solanaceae plant (or ahomolog thereof) containing a mutant version of the gene. In someembodiments, mutations are introduced into two or all three of aSolyc04g005320 gene (or a homolog thereof), a Solyc12g038510 gene (or ahomolog thereof), and a Solyc03g114840 gene (or a homolog thereof).

In any of the methods described herein, the mutant gene can beintroduced into a Solanaceae plant or a plant part or produced in aSolanaceae plant or plant part by any method described herein or knownto those of skill in the art, such as Agrobacterium-mediatedrecombination, viral-vector mediated recombination, microinjection, genegun bombardment/biolistic particle delivery, electroporation,mutagenesis (e.g., by ethyl methanesulfonate or fast neutronirradiation), TILLING (Targeting Induced Local Lesions in Genomes),conventional marker-assisted introgression, and nuclease mediatedrecombination (e.g., use of custom-made restriction enzymes fortargeting mutagenesis by gene replacement, see, e.g., CRISPR-Cas9:Genome engineering using the CRISPR-Cas9 system. Ran F A, Hsu P D,Wright J, Agarwala V, Scott D A, Zhang F. Nat Protoc. 2013 November;8(11):2281-308; TALEN endonucleases: Nucleic Acids Res. 2011 July;39(12):e82. Efficient design and assembly of custom TALEN and other TALeffector-based constructs for DNA targeting. Cermak T, Doyle E L,Christian M, Wang L, Zhang Y, Schmidt C, Baller J A, Somia N V,Bogdanove A J, Voytas D F and Plant Biotechnol J. 2012 May;10(4):373-89. Genome modifications in plant cells by custom-maderestriction enzymes. Tzfira T, Weinthal D, Marton I, Zeevi V, Zuker A,Vainstein A.). Genetically-altered Solanaceae plants produced by orproducible by a method described herein are also claimed.

In some embodiments, the mutation produces a null allele, a hypomorphicallele, or a hypermorphic allele of a Solyc04g005320 gene (or a homologthereof), a Solyc12g038510 gene (or a homolog thereof), or aSolyc03g114840 gene (or a homolog thereof) as described herein. In someembodiments, the mutation is a null mutation of a Solyc04g005320 gene(or a homolog thereof), a Solyc12g038510 gene (or a homolog thereof), ora Solyc03g114840 gene (or a homolog thereof) that is introduced usingCRISPR/Cas9.

Alternatively, a method of producing a genetically-altered Solanaceaeplant comprises a reducing (partially or completely) function of awild-type Solyc04g005320 gene (or a homolog thereof), a wild-typeSolyc12g038510 gene (or a homolog thereof), or a wild-typeSolyc03g114840 gene (or a homolog thereof) in the plant or plant part.In some embodiments, reducing the function comprises performing any ofthe following methods of RNA-interference (e.g., administering to theSolanaceae plant a micro-RNA or a small interfering (si)-RNA or hairpinRNA) or translational blocking (e.g., administering to the Solanaceaeplant a morpholino). Methods of RNA-interference and translationalblocking are well-known in the art. Methods of producing micro-RNAs,si-RNAs, and morpholinos are well-known in the art and can involve useof the nucleotides sequences provided herein.

In some embodiments, the method comprises crossing a producedgenetically-altered Solanaceae plant containing a mutant Solyc04g005320gene (or a homolog thereof) to another genetically-altered Solanaceaeplant comprising a mutant Solyc12g038510 gene (or a homolog thereof), amutant Solyc03g114840 gene (or a homolog thereof), or both a mutantSolyc12g038510 gene (or a homolog thereof) and a mutant Solyc03g114840gene (or a homolog thereof). In some embodiments, the method comprisescrossing a produced genetically-altered Solanaceae plant containing amutant Solyc12g038510 gene (or a homolog thereof) to anothergenetically-altered Solanaceae plant comprising a mutant Solyc04g005320gene (or a homolog thereof), a mutant Solyc03g114840 gene (or a homologthereof), or both a mutant Solyc04g005320 gene (or a homolog thereof)and a mutant Solyc03g114840 gene (or a homolog thereof). In someembodiments, the method comprises crossing a producedgenetically-altered Solanaceae plant containing a mutant Solyc03g114840gene (or a homolog thereof) to another genetically-altered Solanaceaeplant comprising a mutant Solyc12g038510 gene (or a homolog thereof), amutant Solyc04g005320 gene (or a homolog thereof), or both a mutantSolyc12g038510 gene (or a homolog thereof) and a mutant Solyc04g005320gene (or a homolog thereof).

Example Nucleic Acid Sequences of the Disclosure

Wild-type Solyc04g005320 gene (SEQ ID NO: 1)ATGGGAAGAGGTAAGGTAGAATTGAAGAGAATAGAAAATAAGATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGATTACTCAAAAAAGCTTATGAGCTTTCTATTTTGTGTGAAGCTGAAGTTGCTCTTATCATTTTCTCTAATAGAGGCAAACTCTATGAATTTTGCAGTACCTCTAGGTAATATTTTTATGTTTATGTCGTTCCGTTTAAGCTTTACATTTACGTTTTTATACGCAAAACTTTAAATTAGTTCTAAATGTATTAAAAAATTGAAATTTTGAGATTTAATTTCAAAATCTATGGTTAAACGAATGTTTATATGCATTATGATTTTGTTATCTTCTTTTTTTTTAAAAAAAGAAATAAAATATATTGATGTTATAGATCTGAGTGAGAATAGAGTTTTTGGTACATTTATTAAGGGTGAATAATCAAATGTTTCATTTGATTAGATCTAGGTTTTCTTGAACATTAAAATTGTTAAAAAAATTAGTTCATTTTATGAGGTAAATTTTGTTATGATTTGATGTTCCACCTCCATTTTTTCTTATTTTTATTATAAATAAATAAGTTTTAAAATATCCTTACTTTTATATGTTCTTTTAAGTACAGACACATGAATCAAAAAGAAGTTTTATAATATGAATTGAATTAAAGCTGGTTGAATTTCTATCTTCAGTTTTTGAAAACAACTAAAAACTTTGAAAAGGAATTTGATTTTATTATTTATGGCAACAAATAACACCTAACTACTTATCGAGTCGGAATTGACGATATGAATCCTTTAACTTTTCATTTAAGCTCAATTTATATAGAAAATTCTGTATTGTGGATTGAAGTAATTTCTGGAGTTGATCAATTCTATTTAAAAAATTATTTAATTAATAATCATTATCCCAAAAAATTATATTGAAATTAAAAAATAATATTAATTTTTTTAAATAACAAACTTATTAATTGAGTGACCATCTAAATCGTCTTTTTCTTAAAGTTAGGGTCTTGCCTTTCATCTAATTTTGATAGTAATGTTCTTGAACCGACAAATTTTGTCATTTACTCTTATCTGTTATAATTTATGTGATTCGAGTTTTACGAATCAATTTTTGTTTATAATTTCAATCATGTATAAGAAGTATTTTAAGTTATAATAATTAACAATTTTAAGAAAGCATAATCAAGATCAAATAACTTAGTAGAAATAATATTGGTTTATGTAACCTCTATGCATTGACAATATAGTGTTTTTTTTATACTATCAAGTCATTTATTGGATAATTATAATTAAAGAATATTAACTAATGAGTAAATCAATAGTTTAATATTAATGAGTTATCATAGTAGCGTATACTTATTACTCGATATTTGTAATCTAAACATTTTCAATATGCTTAAACTTGATTTTTTTATTTGGATCAAGTATACAATTTTTTTGTTAATAATAAATGACATTGAAACTTATAACTAATTTTATTTAAACAATTTTCTTTCTTTCTTTCCTCAAGGAGAGCATAGTTCTAATTATTATCAATATCATTATTATTATTATCTCTATGTTTATTTTATTATTACTGTTGTTTCTTTTACTTGGATTGTCTGTACTATTTTTACTTCATGGACTTTAATTTTTTGTCTATCGTATTTTTATCATAGTTTTTACTCTTGTATTGGCTAAACCTAGTTTTGAAATTGTTTTTCATAAGCTGAAAGAGTCTATCAAAAACAACTTCTCACGAGATAGAAATAAAGTTTACGTATATTCCATTCTTCTCAAACCCCACTTATGAGATTATACTGAAATGTTACTATTATTATTATACTTTGTAACATGCTAAAAAAACTAGTAATAATTACACTTCTTGCCAAAGAGTAAATAAAGTATGATCCTTTAATAAGTTGAAAATCCCTCTAAATCAAATTATCACTTTTGTGCAACTTGTCTTCTTTTTTTTCTTCTAGTATGTCTGATACACTGGAGAGATACCATAGATGCAGCTATGGTGACCTTGAAACTGGCCAGTCTTCAAAGGATTCACAGGTTACTTCATCTTCCTCAGAATTACAATTTACTAATAAATTTAACTTATATACTCTGACACAGTATCGATGCAATTTAAACCTTTTATAACAGATTATCTGTTTTTATTTTAATTTCTTCGTAAATAATTAATAAGTCGATATTGATAACTAACGCCAAGCACCCTATCTTCATCTAACTAATTAGTGTTATTATGCAATAGAATAACTACCAAGAGTATATGAAGCTGAAAGCAAGAGTTGAAGTGCTACAACAGTCACAAAGGTGATACATTATTTGTTTTAAAAACACTTTTACTTGTCTCATTTTGATTGGCTCATCTGAACACCTGAACCGGTCTAGAAGTATTTTGAACATGCATAATTGGACATGTTCAATCATGCGTTTGTTTGATCAGGTTCAGGATGTTTAGATGAGACCTCGTAAAATAAATTAAGGGGAGGCTTTTTAATATGATATTTGTGTCTCAAATATATCACTTTTCTACCCTAATTCTTAATAACATTGTATTTACTTAATTATTCTTAATCTTTAAGGCATATACTTGGAGAGGACTTAGGACAATTAAACACAAAAGATTTGGAACAGCTTGAGCGTCAACTGGATTCATCTTTGAGGCTAATAAGATCAAGAAGGGTATGTTCTATGCACCTTCAATTTATTTGTCAAATTTTAGGCTTTCAGATCATGTCTTAATCTTAATGTCCGATGACAGTTTCAGTGGCGGAATTAGAAATTTATGCAAGACAATTCAAGCAATATTATATATTATAGAATGTCAGACTTGAAATTTGAACTTGAGACATTGAACCTCTTTACAAATACACTAATATCTAACCTCGTATCAACGGGGTTCAACAATTTATATATATATAAAAAACACTTAATTTTGCCCTATTTGGTGTAATATATAATTTTATCAAAGGTATGTTGGGAAAATGATAAAAATTACTTATGAATAATATCCAAATGGAATAATATAATAACAATTACTTACTATTACTTGATAGTGCCACAAAACTACTAAACCTTAAAATAAGTTCTTTTATTTTACATAATTCATTATAATCTTTGGCATGAATTTACTCAAGCATTGCTTCAGAATGATCAAAGCCTCCTTAATATTTTTGGGTACAGACATAAAGTCTAGACATGCAATCAAAGATATAGATGCACGAGATGACTAATCAAAGGAAACAATAGGAACGATCAAAAAAATTGAAATTGAAAATATATTTTTTTTAAACTAAAGGTAAGTCAAGATTACCAAGTAAGTGTATTATTGTAACTTTTGTATTATTTATCCTAAGTAAACATGTATCAAAAACATACACAAATTTACTTTCTCTTTTATTACTAACATCAACTTACATGCTAATTATAAATAATTAAAGGGTAAATAGTTGGTTGCATGATTTGGTAAAAGAAGTTGTTAACCTACTCTTTGATAACATATATGTTTTCAGACACAAAACATGCTTGATCAACTTTCTGATCTTCAACAAAAGGTATGTATTGTATAATATAATCCCTTAAGTTGACAATTAAATAGATTGTTCAATTGTTAATTTGACATTGTATGTGTTCTTTTTTCTTTTTTTCTACAGGAACAATCTCTTCTTGAAATCAACAGATCCTTGAAAACAAAGGTACAAAGCACACATTTTGGACCTTTTATGAGTTTTTTAGGGCGTGTTTGATTTATTTATTTTTTCTGAATTTTTTCATGTTTGGTTGATCTAAATTCTGGGAAAATACTTTTTTCTATGAAAGTAAGTTTTTTAAAAATGACTTAGCCAGTGGAAGTAGGGAAAACAAGTTGTGACGACATTCCACGTTGATTGTTTTCTCTCGATCTTCCTACACACCTTAAGTTCGCCACCACCTCTCGCAGTATTTGTTTAGATTATATAAAAATGTATCAAGAATGACACTTTTTATTTGTGTACATAATAAAAGAAAATAAGTAAGAAACCGAACATTTTCCCATGGAAAATATTATTTTTCATACCAAACACACCCTTAGTCTTTGTTTTAGGGTATATGACTAATTTGTTCTCCATTTCGGATATTTAGATTCGTATGGGTTTTTCTTTGATGTCTAACTTATCGTACTTTTTACGCGATTTTATGAAATTCTTATAATAGTTGGAAGAAAACTCTGTAGCACATTGGCATATCACTGGAGAGCAAAATGTACAATTCAGACAACAACCTGCTCAGTCAGAGGGGTTCTTTCAGCCTTTACAATGCAATACTAATATAGTGCCAAACAGGTAACATATAATTTTATGTTTTCTTTTTTTCCTTTAAATAGCATATTTTTTGCAACATTTTAAATTGAACCGTTGAATTGAGTCGTTGAGAGGTGAATTCAGAATCTGAAGTAACAGATACATTCAAATTAATTTCTTTGTGTATTTATCGAAGTGAGTCAAGTCGTAAGTCTAGAGGTGAATTTAGAATCTAAAGTAATAGATACAGTCGAATCAATCTCTTTGTGTATTTATTGAAGTGAGTCGTTTTGTAAAGTTTGAGACGAATTCAAAACCTAAAGTAATACTAAATACATACATTCAAAATAATTTCTAAAGCGAGTTATGTTGGAAGTCGAGAGACGAAAGTATATTATATATGGATCAATTCAAATTAATTTCTTAATGTATTTGATGAGCGTTGTTGTAGGGGCGAATTCAGAATCTGAAGTTCATGTAAGTACAGGTACAATGTGGCTCCATTGGATAGTATAGAACCATCAACACAGAATGCTACTGGAATTTTACCAGGATGGATGCTTTGA Wild-type Solyc04g005320 coding sequence (SEQ ID NO: 2)ATGGGAAGAGGTAAGGTAGAATTGAAGAGAATAGAAAATAAGATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGATTACTCAAAAAAGCTTATGAGCTTTCTATTTTGTGTGAAGCTGAAGTTGCTCTTATCATTTTCTCTAATAGAGGCAAACTCTATGAATTTTGCAGTACCTCTAGTATGTCTGATACACTGGAGAGATACCATAGATGCAGCTATGGTGACCTTGAAACTGGCCAGTCTTCAAAGGATTCACAGAATAACTACCAAGAGTATATGAAGCTGAAAGCAAGAGTTGAAGTGCTACAACAGTCACAAAGGCATATACTTGGAGAGGACTTAGGACAATTAAACACAAAAGATTTGGAACAGCTTGAGCGTCAACTGGATTCATCTTTGAGGCTAATAAGATCAAGAAGGACACAAAACATGCTTGATCAACTTTCTGATCTTCAACAAAAGGAACAATCTCTTCTTGAAATCAACAGATCCTTGAAAACAAAGTTGGAAGAAAACTCTGTAGCACATTGGCATATCACTGGAGAGCAAAATGTACAATTCAGACAACAACCTGCTCAGTCAGAGGGGTTCTTTCAGCCTTTACAATGCAATACTAATATAGTGCCAAACAGGTACAATGTGGCTCCATTGGATAGTATAGAACCATCAACACAGAATGCTACTGGAATTTTACCAGGATGGATGCTTTGAMutant Solyc04g005320 gene allele lin^(trans) (SEQ ID NO: 3)ATGGGAAGAGGTAAGGTAGAATTGAAGAGAATAGAAAATAAGATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGATTACTCAAAAAAGCTTATGAGCTTTCTATTTTGTGTGAAGCTGAAGTTGCTCTTATCATTTTCTCTAATAGAGGCAAACTCTATGAATTTTGCAGTACCTCTAGGTAATATTTTTATGTTTATGTCGTTCCGTTTAAGCTTTACATTTACGTTTTTATACGCAAAACTTTAAATTAGTTCTAAATGTATTAAAAAATTGAAATTTTGAGATTTAATTTCAAAATCTATGGTTAAACGAATGTTTATATGCATTATGATTTTGTTATCTTCTTTTTTTTTAAAAAAAGAAATAAAATATATTGATGTTATAGATCTGAGTGAGAATAGAGTTTTTGGTACATTTATTAAGGGTGAATAATCAAATGTTTCATTTGATTAGATCTAGGTTTTCTTGAACATTAAAATTGTTAAAAAAATT

TTCATTTTATGAGGTAAATTTTGTTATGATTTGATGTTCCACCTCCATTTTTTCTTATTTTTATTATAAATAAATAAGTTTTAAAATATCCTTACTTTTATATGTTCTTTTAAGTACAGACACATGAATCAAAAAGAAGTTTTATAATATGAATTGAATTAAAGCTGGTTGAATTTCTATCTTCAGTTTTTGAAAACAACTAAAAACTTTGAAAAGGAATTTGATTTTATTATTTATGGCAACAAATAACACCTAACTACTTATCGAGTCGGAATTGACGATATGAATCCTTTAACTTTTCATTTAAGCTCAATTTATATAGAAAATTCTGTATTGTGGATTGAAGTAATTTCTGGAGTTGATCAATTCTATTTAAAAAATTATTTAATTAATAATCATTATCCCAAAAAATTATATTGAAATTAAAAAATAATATTAATTTTTTTAAATAACAAACTTATTAATTGAGTGACCATCTAAATCGTCTTTTTCTTAAAGTTAGGGTCTTGCCTTTCATCTAATTTTGATAGTAATGTTCTTGAACCGACAAATTTTGTCATTTACTCTTATCTGTTATAATTTATGTGATTCGAGTTTTACGAATCAATTTTTGTTTATAATTTCAATCATGTATAAGAAGTATTTTAAGTTATAATAATTAACAATTTTAAGAAAGCATAATCAAGATCAAATAACTTAGTAGAAATAATATTGGTTTATGTAACCTCTATGCATTGACAATATAGTGTTTTTTTTATACTATCAAGTCATTTATTGGATAATTATAATTAAAGAATATTAACTAATGAGTAAATCAATAGTTTAATATTAATGAGTTATCATAGTAGCGTATACTTATTACTCGATATTTGTAATCTAAACATTTTCAATATGCTTAAACTTGATTTTTTTATTTGGATCAAGTATACAATTTTTTTGTTAATAATAAATGACATTGAAACTTATAACTAATTTTATTTAAACAATTTTCTTTCTTTCTTTCCTCAAGGAGAGCATAGTTCTAATTATTATCAATATCATTATTATTATTATCTCTATGTTTATTTTATTATTACTGTTGTTTCTTTTACTTGGATTGTCTGTACTATTTTTACTTCATGGACTTTAATTTTTTGTCTATCGTATTTTTATCATAGTTTTTACTCTTGTATTGGCTAAACCTAGTTTTGAAATTGTTTTTCATAAGCTGAAAGAGTCTATCAAAAACAACTTCTCACGAGATAGAAATAAAGTTTACGTATATTCCATTCTTCTCAAACCCCACTTATGAGATTATACTGAAATGTTACTATTATTATTATACTTTGTAACATGCTAAAAAAACTAGTAATAATTACACTTCTTGCCAAAGAGTAAATAAAGTATGATCCTTTAATAAGTTGAAAATCCCTCTAAATCAAATTATCACTTTTGTGCAACTTGTCTTCTTTTTTTTCTTCTAGTATGTCTGATACACTGGAGAGATACCATAGATGCAGCTATGGTGACCTTGAAACTGGCCAGTCTTCAAAGGATTCACAGGTTACTTCATCTTCCTCAGAATTACAATTTACTAATAAATTTAACTTATATACTCTGACACAGTATCGATGCAATTTAAACCTTTTATAACAGATTATCTGTTTTTATTTTAATTTCTTCGTAAATAATTAATAAGTCGATATTGATAACTAACGCCAAGCACCCTATCTTCATCTAACTAATTAGTGTTATTATGCAATAGAATAACTACCAAGAGTATATGAAGCTGAAAGCAAGAGTTGAAGTGCTACAACAGTCACAAAGGTGATACATTATTTGTTTTAAAAACACTTTTACTTGTCTCATTTTGATTGGCTCATCTGAACACCTGAACCGGTCTAGAAGTATTTTGAACATGCATAATTGGACATGTTCAATCATGCGTTTGTTTGATCAGGTTCAGGATGTTTAGATGAGACCTCGTAAAATAAATTAAGGGGAGGCTTTTTAATATGATATTTGTGTCTCAAATATATCACTTTTCTACCCTAATTCTTAATAACATTGTATTTACTTAATTATTCTTAATCTTTAAGGCATATACTTGGAGAGGACTTAGGACAATTAAACACAAAAGATTTGGAACAGCTTGAGCGTCAACTGGATTCATCTTTGAGGCTAATAAGATCAAGAAGGGTATGTTCTATGCACCTTCAATTTATTTGTCAAATTTTAGGCTTTCAGATCATGTCTTAATCTTAATGTCCGATGACAGTTTCAGTGGCGGAATTAGAAATTTATGCAAGACAATTCAAGCAATATTATATATTATAGAATGTCAGACTTGAAATTTGAACTTGAGACATTGAACCTCTTTACAAATACACTAATATCTAACCTCGTATCAACGGGGTTCAACAATTTATATATATATAAAAAACACTTAATTTTGCCCTATTTGGTGTAATATATAATTTTATCAAAGGTATGTTGGGAAAATGATAAAAATTACTTATGAATAATATCCAAATGGAATAATATAATAACAATTACTTACTATTACTTGATAGTGCCACAAAACTACTAAACCTTAAAATAAGTTCTTTTATTTTACATAATTCATTATAATCTTTGGCATGAATTTACTCAAGCATTGCTTCAGAATGATCAAAGCCTCCTTAATATTTTTGGGTACAGACATAAAGTCTAGACATGCAATCAAAGATATAGATGCACGAGATGACTAATCAAAGGAAACAATAGGAACGATCAAAAAAATTGAAATTGAAAATATATTTTTTTTAAACTAAAGGTAAGTCAAGATTACCAAGTAAGTGTATTATTGTAACTTTTGTATTATTTATCCTAAGTAAACATGTATCAAAAACATACACAAATTTACTTTCTCTTTTATTACTAACATCAACTTACATGCTAATTATAAATAATTAAAGGGTAAATAGTTGGTTGCATGATTTGGTAAAAGAAGTTGTTAACCTACTCTTTGATAACATATATGTTTTCAGACACAAAACATGCTTGATCAACTTTCTGATCTTCAACAAAAGGTATGTATTGTATAATATAATCCCTTAAGTTGACAATTAAATAGATTGTTCAATTGTTAATTTGACATTGTATGTGTTCTTTTTTCTTTTTTTCTACAGGAACAATCTCTTCTTGAAATCAACAGATCCTTGAAAACAAAGGTACAAAGCACACATTTTGGACCTTTTATGAGTTTTTTAGGGCGTGTTTGATTTATTTATTTTTTCTGAATTTTTTCATGTTTGGTTGATCTAAATTCTGGGAAAATACTTTTTTCTATGAAAGTAAGTTTTTTAAAAATGACTTAGCCAGTGGAAGTAGGGAAAACAAGTTGTGACGACATTCCACGTTGATTGTTTTCTCTCGATCTTCCTACACACCTTAAGTTCGCCACCACCTCTCGCAGTATTTGTTTAGATTATATAAAAATGTATCAAGAATGACACTTTTTATTTGTGTACATAATAAAAGAAAATAAGTAAGAAACCGAACATTTTCCCATGGAAAATATTATTTTTCATACCAAACACACCCTTAGTCTTTGTTTTAGGGTATATGACTAATTTGTTCTCCATTTCGGATATTTAGATTCGTATGGGTTTTTCTTTGATGTCTAACTTATCGTACTTTTTACGCGATTTTATGAAATTCTTATAATAGTTGGAAGAAAACTCTGTAGCACATTGGCATATCACTGGAGAGCAAAATGTACAATTCAGACAACAACCTGCTCAGTCAGAGGGGTTCTTTCAGCCTTTACAATGCAATACTAATATAGTGCCAAACAGGTAACATATAATTTTATGTTTTCTTTTTTTCCTTTAAATAGCATATTTTTTGCAACATTTTAAATTGAACCGTTGAATTGAGTCGTTGAGAGGTGAATTCAGAATCTGAAGTAACAGATACATTCAAATTAATTTCTTTGTGTATTTATCGAAGTGAGTCAAGTCGTAAGTCTAGAGGTGAATTTAGAATCTAAAGTAATAGATACAGTCGAATCAATCTCTTTGTGTATTTATTGAAGTGAGTCGTTTTGTAAAGTTTGAGACGAATTCAAAACCTAAAGTAATACTAAATACATACATTCAAAATAATTTCTAAAGCGAGTTATGTTGGAAGTCGAGAGACGAAAGTATATTATATATGGATCAATTCAAATTAATTTCTTAATGTATTTGATGAGCGTTGTTGTAGGGGCGAATTCAGAATCTGAAGTTCATGTAAGTACAGGTACAATGTGGCTCCATTGGATAGTATAGAACCATCAACACAGAATGCTACTGGAATTTTACCAGGATGGATGCTTTGAMutant Solyc04g005320 gene allele lin^(CR) >allele-1 (SEQ ID NO: 4)ATGGGAAGAGGTAAGGTAGAATTGAAGAGAATAGAAAATAAGATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGATTACTCAAAAAAGCTTATGAGCTTTCTATTTTGTGTGAAGCTGAAGTTGCTCTTATCATTTTCTCTAATAGAGGCAAACTCTATGAATTTTGCAGTACCTCTAGGTAATATTTTTATGTTTATGTCGTTCCGTTTAAGCTTTACATTTACGTTTTTATACGCAAAACTTTAAATTAGTTCTAAATGTATTAAAAAATTGAAATTTTGAGATTTAATTTCAAAATCTATGGTTAAACGAATGTTTATATGCATTATGATTTTGTTATCTTCTTTTTTTTTAAAAAAAGAAATAAAATATATTGATGTTATAGATCTGAGTGAGAATAGAGTTTTTGGTACATTTATTAAGGGTGAATAATCAAATGTTTCATTTGATTAGATCTAGGTTTTCTTGAACATTAAAATTGTTAAAAAAATTAGTTCATTTTATGAGGTAAATTTTGTTATGATTTGATGTTCCACCTCCATTTTTTCTTATTTTTATTATAAATAAATAAGTTTTAAAATATCCTTACTTTTATATGTTCTTTTAAGTACAGACACATGAATCAAAAAGAAGTTTTATAATATGAATTGAATTAAAGCTGGTTGAATTTCTATCTTCAGTTTTTGAAAACAACTAAAAACTTTGAAAAGGAATTTGATTTTATTATTTATGGCAACAAATAACACCTAACTACTTATCGAGTCGGAATTGACGATATGAATCCTTTAACTTTTCATTTAAGCTCAATTTATATAGAAAATTCTGTATTGTGGATTGAAGTAATTTCTGGAGTTGATCAATTCTATTTAAAAAATTATTTAATTAATAATCATTATCCCAAAAAATTATATTGAAATTAAAAAATAATATTAATTTTTTTAAATAACAAACTTATTAATTGAGTGACCATCTAAATCGTCTTTTTCTTAAAGTTAGGGTCTTGCCTTTCATCTAATTTTGATAGTAATGTTCTTGAACCGACAAATTTTGTCATTTACTCTTATCTGTTATAATTTATGTGATTCGAGTTTTACGAATCAATTTTTGTTTATAATTTCAATCATGTATAAGAAGTATTTTAAGTTATAATAATTAACAATTTTAAGAAAGCATAATCAAGATCAAATAACTTAGTAGAAATAATATTGGTTTATGTAACCTCTATGCATTGACAATATAGTGTTTTTTTTATACTATCAAGTCATTTATTGGATAATTATAATTAAAGAATATTAACTAATGAGTAAATCAATAGTTTAATATTAATGAGTTATCATAGTAGCGTATACTTATTACTCGATATTTGTAATCTAAACATTTTCAATATGCTTAAACTTGATTTTTTTATTTGGATCAAGTATACAATTTTTTTGTTAATAATAAATGACATTGAAACTTATAACTAATTTTATTTAAACAATTTTCTTTCTTTCTTTCCTCAAGGAGAGCATAGTTCTAATTATTATCAATATCATTATTATTATTATCTCTATGTTTATTTTATTATTACTGTTGTTTCTTTTACTTGGATTGTCTGTACTATTTTTACTTCATGGACTTTAATTTTTTGTCTATCGTATTTTTATCATAGTTTTTACTCTTGTATTGGCTAAACCTAGTTTTGAAATTGTTTTTCATAAGCTGAAAGAGTCTATCAAAAACAACTTCTCACGAGATAGAAATAAAGTTTACGTATATTCCATTCTTCTCAAACCCCACTTATGAGATTATACTGAAATGTTACTATTATTATTATACTTTGTAACATGCTAAAAAAACTAGTAATAATTACACTTCTTGCCAAAGAGTAAATAAAGTATGATCCTTTAATAAGTTGAAAATCCCTCTAAATCAAATTATCACTTTTGTGCAACTTGTCTTCTTTTTTTTCTTCTAGTATGTCCCATAGATGCAGCTATGGTGACCTTGAAACTGGCCAGTCTTCAAAGGATTCACAGGTTACTTCATCTTCCTCAGAATTACAATTTACTAATAAATTTAACTTATATACTCTGACACAGTATCGATGCAATTTAAACCTTTTATAACAGATTATCTGTTTTTATTTTAATTTCTTCGTAAATAATTAATAAGTCGATATTGATAACTAACGCCAAGCACCCTATCTTCATCTAACTAATTAGTGTTATTATGCAATAGAATAACTACCAAGAGTATATGAAGCTGAAAGCAAGAGTTGAAGTGCTACAACAGTCACAAAGGTGATACATTATTTGTTTTAAAAACACTTTTACTTGTCTCATTTTGATTGGCTCATCTGAACACCTGAACCGGTCTAGAAGTATTTTGAACATGCATAATTGGACATGTTCAATCATGCGTTTGTTTGATCAGGTTCAGGATGTTTAGATGAGACCTCGTAAAATAAATTAAGGGGAGGCTTTTTAATATGATATTTGTGTCTCAAATATATCACTTTTCTACCCTAATTCTTAATAACATTGTATTTACTTAATTATTCTTAATCTTTAAGGCATATACTTGGAGAGGACTTAGGACAATTAAACACAAAAGATTTGGAACAGCTTGAGCAACTGGATTCATCTTTGAGGCTAATAAGATCAAGAAGGGTATGTTCTATGCACCTTCAATTTATTTGTCAAATTTTAGGCTTTCAGATCATGTCTTAATCTTAATGTCCGATGACAGTTTCAGTGGCGGAATTAGAAATTTATGCAAGACAATTCAAGCAATATTATATATTATAGAATGTCAGACTTGAAATTTGAACTTGAGACATTGAACCTCTTTACAAATACACTAATATCTAACCTCGTATCAACGGGGTTCAACAATTTATATATATATAAAAAACACTTAATTTTGCCCTATTTGGTGTAATATATAATTTTATCAAAGGTATGTTGGGAAAATGATAAAAATTACTTATGAATAATATCCAAATGGAATAATATAATAACAATTACTTACTATTACTTGATAGTGCCACAAAACTACTAAACCTTAAAATAAGTTCTTTTATTTTACATAATTCATTATAATCTTTGGCATGAATTTACTCAAGCATTGCTTCAGAATGATCAAAGCCTCCTTAATATTTTTGGGTACAGACATAAAGTCTAGACATGCAATCAAAGATATAGATGCACGAGATGACTAATCAAAGGAAACAATAGGAACGATCAAAAAAATTGAAATTGAAAATATATTTTTTTTAAACTAAAGGTAAGTCAAGATTACCAAGTAAGTGTATTATTGTAACTTTTGTATTATTTATCCTAAGTAAACATGTATCAAAAACATACACAAATTTACTTTCTCTTTTATTACTAACATCAACTTACATGCTAATTATAAATAATTAAAGGGTAAATAGTTGGTTGCATGATTTGGTAAAAGAAGTTGTTAACCTACTCTTTGATAACATATATGTTTTCAGACACAAAACATGCTTGATCAACTTTCTGATCTTCAACAAAAGGTATGTATTGTATAATATAATCCCTTAAGTTGACAATTAAATAGATTGTTCAATTGTTAATTTGACATTGTATGTGTTCTTTTTTCTTTTTTTCTACAGGAACAATCTCTTCTTGAAATCAACAGATCCTTGAAAACAAAGGTACAAAGCACACATTTTGGACCTTTTATGAGTTTTTTAGGGCGTGTTTGATTTATTTATTTTTTCTGAATTTTTTCATGTTTGGTTGATCTAAATTCTGGGAAAATACTTTTTTCTATGAAAGTAAGTTTTTTAAAAATGACTTAGCCAGTGGAAGTAGGGAAAACAAGTTGTGACGACATTCCACGTTGATTGTTTTCTCTCGATCTTCCTACACACCTTAAGTTCGCCACCACCTCTCGCAGTATTTGTTTAGATTATATAAAAATGTATCAAGAATGACACTTTTTATTTGTGTACATAATAAAAGAAAATAAGTAAGAAACCGAACATTTTCCCATGGAAAATATTATTTTTCATACCAAACACACCCTTAGTCTTTGTTTTAGGGTATATGACTAATTTGTTCTCCATTTCGGATATTTAGATTCGTATGGGTTTTTCTTTGATGTCTAACTTATCGTACTTTTTACGCGATTTTATGAAATTCTTATAATAGTTGGAAGAAAACTCTGTAGCACATTGGCATATCACTGGAGAGCAAAATGTACAATTCAGACAACAACCTGCTCAGTCAGAGGGGTTCTTTCAGCCTTTACAATGCAATACTAATATAGTGCCAAACAGGTAACATATAATTTTATGTTTTCTTTTTTTCCTTTAAATAGCATATTTTTTGCAACATTTTAAATTGAACCGTTGAATTGAGTCGTTGAGAGGTGAATTCAGAATCTGAAGTAACAGATACATTCAAATTAATTTCTTTGTGTATTTATCGAAGTGAGTCAAGTCGTAAGTCTAGAGGTGAATTTAGAATCTAAAGTAATAGATACAGTCGAATCAATCTCTTTGTGTATTTATTGAAGTGAGTCGTTTTGTAAAGTTTGAGACGAATTCAAAACCTAAAGTAATACTAAATACATACATTCAAAATAATTTCTAAAGCGAGTTATGTTGGAAGTCGAGAGACGAAAGTATATTATATATGGATCAATTCAAATTAATTTCTTAATGTATTTGATGAGCGTTGTTGTAGGGGCGAATTCAGAATCTGAAGTTCATGTAAGTACAGGTACAATGTGGCTCCATTGGATAGTATAGAACCATCAACACAGAATGCTACTGGAATTTTACCAGGATGGATGCTTTGA >allele-2 (SEQ ID NO: 5)ATGGGAAGAGGTAAGGTAGAATTGAAGAGAATAGAAAATAAGATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGATTACTCAAAAAAGCTTATGAGCTTTCTATTTTGTGTGAAGCTGAAGTTGCTCTTATCATTTTCTCTAATAGAGGCAAACTCTATGAATTTTGCAGTACCTCTAGGTAATATTTTTATGTTTATGTCGTTCCGTTTAAGCTTTACATTTACGTTTTTATACGCAAAACTTTAAATTAGTTCTAAATGTATTAAAAAATTGAAATTTTGAGATTTAATTTCAAAATCTATGGTTAAACGAATGTTTATATGCATTATGATTTTGTTATCTTCTTTTTTTTTAAAAAAAGAAATAAAATATATTGATGTTATAGATCTGAGTGAGAATAGAGTTTTTGGTACATTTATTAAGGGTGAATAATCAAATGTTTCATTTGATTAGATCTAGGTTTTCTTGAACATTAAAATTGTTAAAAAAATTAGTTCATTTTATGAGGTAAATTTTGTTATGATTTGATGTTCCACCTCCATTTTTTCTTATTTTTATTATAAATAAATAAGTTTTAAAATATCCTTACTTTTATATGTTCTTTTAAGTACAGACACATGAATCAAAAAGAAGTTTTATAATATGAATTGAATTAAAGCTGGTTGAATTTCTATCTTCAGTTTTTGAAAACAACTAAAAACTTTGAAAAGGAATTTGATTTTATTATTTATGGCAACAAATAACACCTAACTACTTATCGAGTCGGAATTGACGATATGAATCCTTTAACTTTTCATTTAAGCTCAATTTATATAGAAAATTCTGTATTGTGGATTGAAGTAATTTCTGGAGTTGATCAATTCTATTTAAAAAATTATTTAATTAATAATCATTATCCCAAAAAATTATATTGAAATTAAAAAATAATATTAATTTTTTTAAATAACAAACTTATTAATTGAGTGACCATCTAAATCGTCTTTTTCTTAAAGTTAGGGTCTTGCCTTTCATCTAATTTTGATAGTAATGTTCTTGAACCGACAAATTTTGTCATTTACTCTTATCTGTTATAATTTATGTGATTCGAGTTTTACGAATCAATTTTTGTTTATAATTTCAATCATGTATAAGAAGTATTTTAAGTTATAATAATTAACAATTTTAAGAAAGCATAATCAAGATCAAATAACTTAGTAGAAATAATATTGGTTTATGTAACCTCTATGCATTGACAATATAGTGTTTTTTTTATACTATCAAGTCATTTATTGGATAATTATAATTAAAGAATATTAACTAATGAGTAAATCAATAGTTTAATATTAATGAGTTATCATAGTAGCGTATACTTATTACTCGATATTTGTAATCTAAACATTTTCAATATGCTTAAACTTGATTTTTTTATTTGGATCAAGTATACAATTTTTTTGTTAATAATAAATGACATTGAAACTTATAACTAATTTTATTTAAACAATTTTCTTTCTTTCTTTCCTCAAGGAGAGCATAGTTCTAATTATTATCAATATCATTATTATTATTATCTCTATGTTTATTTTATTATTACTGTTGTTTCTTTTACTTGGATTGTCTGTACTATTTTTACTTCATGGACTTTAATTTTTTGTCTATCGTATTTTTATCATAGTTTTTACTCTTGTATTGGCTAAACCTAGTTTTGAAATTGTTTTTCATAAGCTGAAAGAGTCTATCAAAAACAACTTCTCACGAGATAGAAATAAAGTTTACGTATATTCCATTCTTCTCAAACCCCACTTATGAGATTATACTGAAATGTTACTATTATTATTATACTTTGTAACATGCTAAAAAAACTAGTAATAATTACACTTCTTGCCAAAGAGTAAATAAAGTATGATCCTTTAATAAGTTGAAAATCCCTCTAAATCAAATTATCACTTTTGTGCAACTTGTCTTCTTTTTTTTCTTCTAGTATGTCTGATACACTGGAGAGATACCATAGATGCAGCTATGGTGACCTTGAAACTGGCCAGTCTTCAAAGGATTCACAGGTTACTTCATCTTCCTCAGAATTACAATTTACTAATAAATTTAACTTATATACTCTGACACAGTATCGATGCAATTTAAACCTTTTATAACAGATTATCTGTTTTTATTTTAATTTCTTCGTAAATAATTAATAAGTCGATATTGATAACTAACGCCAAGCACCCTATCTTCATCTAACTAATTAGTGTTATTATGCAATAGAATAACTACCAAGAGTATATGAAGCTGAAAGCAAGAGTTGAAGTGCTACAACAGTCACAAAGGTGATACATTATTTGTTTTAAAAACACTTTTACTTGTCTCATTTTGATTGGCTCATCTGAACACCTGAACCGGTCTAGAAGTATTTTGAACATGCATAATTGGACATGTTCAATCATGCGTTTGTTTGATCAGGTTCAGGATGTTTAGATGAGACCTCGTAAAATAAATTAAGGGGAGGCTTTTTAATATGATATTTGTGTCTCAAATATATCACTTTTCTACCCTAATTCTTAATAACATTGTATTTACTTAATTATTCTTAATCTTTAAGGCATATACTTGGAGAGGACTTAGGACAATTAAACACAAAAGATTTGGAAAACTGGATTCATCTTTGAGGCTAATAAGATCAAGAAGGGTATGTTCTATGCACCTTCAATTTATTTGTCAAATTTTAGGCTTTCAGATCATGTCTTAATCTTAATGTCCGATGACAGTTTCAGTGGCGGAATTAGAAATTTATGCAAGACAATTCAAGCAATATTATATATTATAGAATGTCAGACTTGAAATTTGAACTTGAGACATTGAACCTCTTTACAAATACACTAATATCTAACCTCGTATCAACGGGGTTCAACAATTTATATATATATAAAAAACACTTAATTTTGCCCTATTTGGTGTAATATATAATTTTATCAAAGGTATGTTGGGAAAATGATAAAAATTACTTATGAATAATATCCAAATGGAATAATATAATAACAATTACTTACTATTACTTGATAGTGCCACAAAACTACTAAACCTTAAAATAAGTTCTTTTATTTTACATAATTCATTATAATCTTTGGCATGAATTTACTCAAGCATTGCTTCAGAATGATCAAAGCCTCCTTAATATTTTTGGGTACAGACATAAAGTCTAGACATGCAATCAAAGATATAGATGCACGAGATGACTAATCAAAGGAAACAATAGGAACGATCAAAAAAATTGAAATTGAAAATATATTTTTTTTAAACTAAAGGTAAGTCAAGATTACCAAGTAAGTGTATTATTGTAACTTTTGTATTATTTATCCTAAGTAAACATGTATCAAAAACATACACAAATTTACTTTCTCTTTTATTACTAACATCAACTTACATGCTAATTATAAATAATTAAAGGGTAAATAGTTGGTTGCATGATTTGGTAAAAGAAGTTGTTAACCTACTCTTTGATAACATATATGTTTTCAGACACAAAACATGCTTGATCAACTTTCTGATCTTCAACAAAAGGTATGTATTGTATAATATAATCCCTTAAGTTGACAATTAAATAGATTGTTCAATTGTTAATTTGACATTGTATGTGTTCTTTTTTCTTTTTTTCTACAGGAACAATCTCTTCTTGAAATCAACAGATCCTTGAAAACAAAGGTACAAAGCACACATTTTGGACCTTTTATGAGTTTTTTAGGGCGTGTTTGATTTATTTATTTTTTCTGAATTTTTTCATGTTTGGTTGATCTAAATTCTGGGAAAATACTTTTTTCTATGAAAGTAAGTTTTTTAAAAATGACTTAGCCAGTGGAAGTAGGGAAAACAAGTTGTGACGACATTCCACGTTGATTGTTTTCTCTCGATCTTCCTACACACCTTAAGTTCGCCACCACCTCTCGCAGTATTTGTTTAGATTATATAAAAATGTATCAAGAATGACACTTTTTATTTGTGTACATAATAAAAGAAAATAAGTAAGAAACCGAACATTTTCCCATGGAAAATATTATTTTTCATACCAAACACACCCTTAGTCTTTGTTTTAGGGTATATGACTAATTTGTTCTCCATTTCGGATATTTAGATTCGTATGGGTTTTTCTTTGATGTCTAACTTATCGTACTTTTTACGCGATTTTATGAAATTCTTATAATAGTTGGAAGAAAACTCTGTAGCACATTGGCATATCACTGGAGAGCAAAATGTACAATTCAGACAACAACCTGCTCAGTCAGAGGGGTTCTTTCAGCCTTTACAATGCAATACTAATATAGTGCCAAACAGGTAACATATAATTTTATGTTTTCTTTTTTTCCTTTAAATAGCATATTTTTTGCAACATTTTAAATTGAACCGTTGAATTGAGTCGTTGAGAGGTGAATTCAGAATCTGAAGTAACAGATACATTCAAATTAATTTCTTTGTGTATTTATCGAAGTGAGTCAAGTCGTAAGTCTAGAGGTGAATTTAGAATCTAAAGTAATAGATACAGTCGAATCAATCTCTTTGTGTATTTATTGAAGTGAGTCGTTTTGTAAAGTTTGAGACGAATTCAAAACCTAAAGTAATACTAAATACATACATTCAAAATAATTTCTAAAGCGAGTTATGTTGGAAGTCGAGAGACGAAAGTATATTATATATGGATCAATTCAAATTAATTTCTTAATGTATTTGATGAGCGTTGTTGTAGGGGCGAATTCAGAATCTGAAGTTCATGTAAGTACAGGTACAATGTGGCTCCATTGGATAGTATAGAACCATCAACACAGAATGCTACTGGAATTTTACCAGGATGGATGCTTTGA Wild-type Solyc12g038510 gene(SEQ ID NO: 6)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGGTATATATATATATACATATGTTTTTCTTCTTTTTGTGTGTGCGTATGTGTTTACTTACTTTCATTAATTAACTCAACCATATATATACATCTCTCACCTCAATTATATATATGTTTGAGATCTGAATGTCTACGGACTCCATTTAGGTACATATCTTTGTTTAGATCATAAATCATCTATCTTCATTCCTAAGATCTACTAATATATATGTATAAGAAGATCCATCCATCTATTAGGTTTTTCAACAACATATACAGTGAAATCTTATATGTGGGCCCACGTATAGCCATATGAGAAAATAGTGTGCACGTAAACATTATCATTACTTAATTATAGGAATATACATCCATTAGGTTTATCAACAACAATAAAATCCTCTAAATGGAGTCTAGTCATAGGTCTAGCCGTTTGAAAATGTAAAATATATGCCGATCTTATCACTATGTCATAATAATAGATATGTTGTTATTGAAAGATTCTCAATCTTTTTTTTTCTTCAAGGTAGAGATTCTTAAGTGGATTCATGTTTTTTTTATCAAAAAAGAAAAAAACAAAAGTGTCCATTTGTTCATCTAATGGGTTTTCCATGTTACCAATTCACTACACTGTTGAGATTTGATTATCAGATGTGTCAAGTTTCGTTTGGTTCCCTAGAAGGGAGAAAAGGCTGCTTATGCAGGCAGGGTATTAAAGATGATATTAATATCTGCAGTAATCAGTAACAGAATATATAAACTTAATAATAAACTTGAAGGTACTTAATTATCCAGCAGATAATCTTCTGTCTCACCGTACACTTTTGTTATATCATAAGCATAAGAATTGTTTTATCAAATATTACCAAACAAAACTTAGTTTTGTTTGGTAATATTTTATAAAATATGTTACCGAAAGTTACTTCCTATAACATATTTTATAAAGAAAAAAATTAAAAACTCCATATACCTAAGAAATGTAACCCCCCCTCCATAACAACAATTTAACAAAAATAAAAACCTACTTTTTTTGAATTTGGTAAATTAGTTTTCTATCCTTTTTAGTAACTTCCTTTCTTATTTTCTTTTTATATTGGTAAAGTTTAATATTACACATTATTTTAACATGTTATAATTTTTTGTGATGCTTAATTATTTGATACATGTAATAAACCATATATTAGAGCTATAAATCAATGACAATGCATGTAGATACAACTCATTTATGATATATTTTGTTTATATATATAACCAATTAGATAATTTGTCTGCGCTTTGTGCAGTCATAAATAATAATTGCATTGAACTTGCAAATATTTTTTTTTAATATCCATACATTAAAAAAAAAGAAAGAGGAAAATTGGTTCCTAAAATATTAGCAATATTCAAACATTTATTTGATTATTAATCATTATCACATAACTTAAGAACGTCTAATGAATGAATTATTCACGAAATAATAAATCATTGGTTCTAAAAAGGAATTTCGTAATAAAATAAAAATTTAAGTTACCATATTCAAAAAAAGAAATTGTGCTTGAACATGAAAATAATTATAATTTTTGAACTTGTATAATGAATTTCTTCAATTCATAAGTGGGAAATTTCATATTTATGTAATAATAGATAATATGTAAGCTCTAATATAGTACTTTAGGTTATAGAATTTAATATAAAATATCAAAACATGAATTCTTGAAATTGAGTAGAGTAATTATTTTCTGCACAATGAATCGGAGACAATAACTTTGAAGAAATATAAACAATAGAGTTCAAAAGATGTAGTCAAAAACAACAATTAATATCATAAGAATAAATTAATGAGTGTAAAAATGCATACCACGATATGTAAAAACAGAATGGAATATAATAAAAAAAATCGAGTTCACTGAATACACAATGTTCCTTTAAGAAAATTATTCTCCTCCAATACCAACGAGATTACATCCTCTAAGGATGGAAATGATTTCATTCCCCAACTTATCCATATAAAAATAGTGGTGTTAGTATGTAACTCAATAGGAGTAAAATACACAAATATTTAATTTTGCGAAAGTAGAAGAAGAAGATCATATTTTTTTTTTAAAATGAGAGGATATATCACTATTTTTAAACAACAAAGGGTAGTGTTAACAAATTTTTATTGTGTCTTGTCTAAAAGGTTACAGCTATTTGAAAAAGTTACAACACTTCGAAAAGTGAACAACATTTCATAAAAGTCGTAACTTTTCATAAAGTCGTAACTCTTCATAAATGTCGCAACTCTTCATAAAAATTACAACTATTGATAAAAGTCACCACTCTTGATAAAGATCACCACTCTTCATTGAAGTTGCAACTTTTCATAAAAATCACATCTTTTAATAAAAAAGAAAGACTAGTTTTTGGAATAAATTAATTTAAAAGAAAATTTTTGTTTGTGGTGGGGCGCCAAGTAGGCAGGCGTAGGGTTCTTTTTATATAAATATATATGATATATGATTCAATATTTGATATATATATATATAGAGAGAGAGATGACAATATAAGACAATTGCAAAAAATAAAATAAAAAACTAATCGAGTAAGTAGGCAAAAAATTATTTATAAAATATATGTAGAATTTCTTTATCAGATATGACTGCCCAAATCTTATATTCAAACTAAAATGCAAGATCAATGGTGCTATATATAGGGTTTTACACAAAAATCAAGATCTAGTCTTGCAAATTTAAATAAAAAACAGTGGTTTACGATGAGATAATGTAGCTTTTGTAAACAATAAAACTAGAAAAATAAATGCAAAGGCATTTTAAAGGATATAATAATGAAGATCAAAGGCAGAGAAGGGAAGAGGCAGCAATATAATGAAGGTAACATCATGGTTCCATTCTAATATATATGCTATTTTTCTTTAGTAAATTTCAAAAATAATGATACATTTTCATATTTGATAAATATTTAATGATACTATCAACATTTTATCTATATTGAGTTCCATTTATTTGACCAAAACCTCACAAAGATGTGCTCTTCGATCTATTCAAAATTTATTCAATTTAAGGATAGCTTTAAAACATGACAAAGTTTTCTCATATATTTCTTAAATTTTATATCCAGTCTAAATACGTATATAAACTAAAATGAAGAGAATAATATGAAGCTTTATTTGATGACATTGTTGAAATAACCAAAAGCTATAAGTGATACAATAGTAAATTTACCATTGGTCAATTCAGAATTATTTAAAAGCTAAAAAAGTCATATAAGTTGGGGTTGCTCAATGTATAGTTTTTGGCTTGTTTTAAGCATTTTAAAACTTTTTTTAAGCGCTTTTTAACATTGCTAAACACTCAAAAAATGATAAATAGTATTTAAATTTGATATGATTAGCTTAAAAGTGAACTCATATACCTTCAAAGTAAAAATCCCCAATTCGAGCTTTCAAACCACTTGATTTTGTGGATGAAATTATACTGAAGTTGAATATATCACTATTTATAGGGGTTAGTGAACTAATACCTTTGATTATTTGGTAGAAATATGTATCTTAGATCACCCTAATGAGCTCCCACTTTTAAAATAGGAAAAACCTCATATGAAGTTCATCACTGTTCATTATATATCACTTTTATTCAAAAACGTTTACAAATGTTCATTGTGACTAAATACCCTTGAGTGTCGAGTTTTCACACCAATAAGGCCTAATTAATAGGTAAACAAAACTATGTCAATCTTCAAAACGCAAATCTAATTATATTTTTAACAAGATTAGAGGTATATATACATATTCTCTTATGTTAACTCTTATTCATTATTGAACAAACTAAGTAAGTGTACCCAAGGTCTCAAACAACAGTTGGTACATTCTTTGTATGTCTTCCTTTGTCTCTTAATAGTCGTCTCCTCCTGTCGATGATTCCTCCAAATACATTAATCAAAGGAAAATCTTTCGCCCTCAACTTGCAAACTTGTCTATCTAAAATTGTTAACAAAGTTTCTTCATTAGAGAAACTATGATTTCTTGAATGTAGCAATTTGATGTGCCATGACTATCATCTTGATCAACATGCTTCTTAACCATCAAAAGATCCTAAACTAGATGCATGTCATGTTAGGAGACATATTAAGCTTGTATATAACTACACCAACATGCTTTAGGATCTCATAAGATCCAAAATTTCTTATTTGGGAGATTTTCAATCCAACAACCATCATAATGAGCAACGTGATGTTATAACATCTCTCTCACACTGCCAGAACAGTCTTATACCTTGTCGGAGTGAAGGACATCCTTAACTAAGTAGATTCACTAAGCTATACTTAAAAAGCAATAAGGAATCATCTAAAATGTGTGACTCTTAACCCATATTGGCATACATGGTTTATGGGGGTTATTAATTGTCTGAACACTCCCCCATATAAATCAGTGATCAATATTAATCCCAATAATATACACTATTATGATTTGAGACTACACCCTGGAAGTGGCCGGCTCTCAAGAACCATTGCTGATCTCCAAGCCAAACCCTCATTCTGGTTGACTACAAGCTGAAGGCAAACTCAAGTATACAAAGCTTAAAACATAATAAAAATAATATACTCAACTCGCCACAAAATAGGCATTTAAGTCTTTAAAACATTTTTAAAAATAAATGAAACAAACTTCTCAAACTGTAATGTATATCTATGAAGCCTCTAAATGAAAAAAATGAAGGCAGATGAGACATACGGCATCCTAACAACTGATATAACTAAGAGTACAAGTGGAGCCCTTCGGATGTAAGGAGGCTCATCAAAGCTAATGTGAACTCCATGTGGTATCAATGAAGCACCTATTGATGACCGTGAATACATGTATCTGCATCATGAAACGATGCAGGCCAAAGGGCTTAGTACGTGAAATGTACGAGCATGTAAAGGGAATTCAAATACATAAACATAGGCTTGAACTTTGATATAAAGGAAACATACTTACCTATTTTTAACTCAAGAATAAAAAACATAGTTCAACTCAATGAAAAGACACTCAAGTCAGTGAAATAGGCCGCAACTCAATAATAAGATATTCGACTATGGGTAATCAACTCTGGGTACTCTATTCAATATAAAGTAAGAATACAAATGCATTATATGGAAAGACTTTAAAACGGTAGAAAACAACTCAATGTATTGAAAATTCAATAGTAAATTAGTTTGTATGTAAGGAACAATATAAACTTTGTTTGTATATGAAAATACAAAATAAACTTTGTGTATATAAAAGTACAAAATATCTCTGTGAAAGTTTCTCTAACCAACAACCATCACTATGAGCTTTCTGATAATACCACGTTTCGCCCATGATGTCAGAACTGTCCTATGATTTTCCAGTTCATAAGACCTACTCACTAAGTGGATCCACAAGTCTATGCTAAAAAATATTTAAGGAATCGTCTAAAAAGTATGACTCATTCTACCCACGTTGGCTACATGATTTATGGGGGTCGTAAGTTATCTAAACTCTCCTCCATATCGATGCGTAATGCTACTCACAAATATACTAGCTCACATGTTTAAAAATATAACTCGTTTTGTTTGAGATCATTACTCAAAATCCTTCTCTTAAAAGAGATGATACTCAAACTGCTCAAAACTCTTTTGGAAATCTCAAATTCGTCTCATCTTAAATGTAAAAATATTTACTCTTGGGAATACATAGTTATCATATATCATTTTAAAGAAAATGAACTCAACTCTGTTCTTTCTCAACTCAAGTGCTCAGTCTTAAACCAAATTAAAAAAAAGACTTCTCAAAATAAAGTTTATGTCGAATTATGGACGTGAACAATTCAATTCAAAGTTTTCGATAACCATAACTAAAACTAAATACTCGAGACTCAACATCTTAGAACTCAAGAACTTAAATGGTAATACTTCTTTCAAGAATGCTCGACTCAGAAGGTTAATGCAGAATAATGTGCATGAATTACTCAACTAAAGGACTCACTGATACTACTCAATCTCAAGATTGCTCGACTCGTAGGGTTAATGCAGAATTATGTGCATGAACTACTCAACTCAAAGACCTTCATAGGTAACATGTAGTAGCCCCATGATTTGGAATATAATCCCAAAATGATTAGGAACTCAATACTCAGGACTTAGAACTTGAAGATAATACTACTTCTCTCAAAGATACCCAACTGACGGAGTTCATGCAGAATTTATGGGCATGAACTACTCGACTCAAGAGTCTAAAACACAATATGACACTCATGTATATAACTCTTCTCATTCTAATACTTGTTTTCTCAAAACTCGGTTTAACTAAATAGTTGATCTCAAAGGATTCACAATTGAACTCAAAGACTTTCTTTGACTCCACTCTTAATTCTCTCTTAAATTTGTATTTGAATTATGAATTTAAGAGTTATGATTCATGATATGGGGAATCTCAATAACAATATAGAAATTTGATAATTAGGAATAGTACTTTTAAAAGAAAACATGAATTCAACTTAAAATCAACTTATCTAAAAAATATTCAAATATAGGGAAAGTATCCTAGACTACTGTGCTACTGATCTGAAAGTAGATGTAGGATGTGAGGATGAACTAGTCCAACACTATGATAGCCTTACATACCTGGAATAACGAGGTTCTTGGAAAATCTTCACTTGAAGAAGAACTTGATTAGAAGCCTTGAAACCTAGCTTGAAGGTAAACAATCAAGAAAACCTTTCTTAAGATTCTTGAATTAGTTTATGAAAATCTCTATGACCAAGCATTTTGATTTTCACTAGTGATTCATAATTGTATGGAGGAATTTGAATTGAAAAAGATGAAATGCTTGGAGAAAAGCTATCTTTGAAGAAGCTTGAAAAAGATTGGAAAGTCCTGTACTTTGATTTTCCCTTAGGATTTTGTCTTAGGGTTTGAGATAGAAAAGAATGATGGACTAAAAGATGAAAATCTAATTGTTTGGATCCTTTTTCAGCCAAGAAATCCGTTTAGGGTTTTCTTGGAGACAAACAAAATAAAAAAGACCATTTTTAATATTTTTCCGTCGGCTAATTCGTAATAACATTGTATCATGTTATTGAAAGAGTCATAACTTTTTACTCAAAAATTGGATTGATGCGAAATTAGTGGTGTTGGAAAGTAGATTCAAGTACCTCTAATTGGATAGGTTATTCCCTACATAAGTCTTTATATTCTAAAAGATATGGTTGTTTGCACTTGACCTAAGTAGAATTTTACATGAAAACTTAATAGAGAAGGAAACTTCAAGAACTCATCAAGAAATTTCAATTGCTCAATATTTATGGATAAATTTGTAGAAGAAACTCATGATTGACATGCGGGTGAATAAACCCAACACTATGGAAGCTTACATACCTCAAAGAACTAGGTTCTTGGCGAAATCTTGAATTTCTTCAACGAACGCTTGAAACTTTGAACTTTTTCTCTTCTTGAACTCTCAACTAAAACCCTAGGCGTATATTAGGATTATAAAAGTTAACATGATAGGATTAGACCTTTAAAAACTTTCTAAAATGAATTAAATCTGATTTAGCATGAAAAAGACCAAAATACCCCTTACTATTTTCGGATAACTTTTCTTAATTGGACTGCCTGACTTCAAAAAGGTATATCTCACTCATCCGACCTCAAAATTTAGCAAATTCAGTGGCGTTAGAAAGCTAATTTAAACACCTTTCATTTTCCATCTCATGGCACACATAACTCATTCTTTAAAGAGAGCTATGATCGTTCAAATTAACTCAAATCTTAGAAGAATTTAGGAATGTCTTGAACGAGCTACATCTAGTGACCTTAACACTTTGGAAAATTTTAAATTTCTTAGTAAAAACTTACTCACTATGAAGGATGGTTCAAGTCTTAGCTCAAAATTTTCCTAAGTTGCTATATATACTCATGCTCATATGTTTAAAACCAAAACCCTTCCTCGATTTGAATTAATTACCAAAAAGATTCTCTTAAAAAGATAATGCTCAAAACTCCCCCTAAACTCATTTGGAAATCTAGGTTTCCCTTGTTTTAAATATAAAAACATTTACTCTTGGAAATATTTAGTTCTCAGATATTCACTTGAAAAAAATTAAACTCGACTCTCATCATCTTCATACTCAAGTGCTCAAGTCCTAAAACAATTTATAACTAATTGTATAAGACTTCTCAAAATAGGGTTCATTCCGAATTATGGACGTGAACGACTCAATTCAAGGATTTCAATAACCATATATATAACTCAATAATAGGAACTCAACAACTCCAGAACTCAATGATACTACTCATCTCAAGAATGCTCGACTCACAGGGTCTTTGCGAAATTATTGGGCATGAACAACTCAACTCAAAGACCTTCATTTATACCATATGGTAGTCCCATAATAGGAATATAATCCCAAAAAAATTAGGAACTCAATACTCAAAAACTTAGAACTCGAAGATATTACTCATCTCAAAGATATTCAATTTATGGAATTCATGCTGAATTATGAGCATGAACGACTTGACTCAAGGATCTCAATAATAATGTAGACTCATGAATACACTCTTCTCATTCTCATACTCACATACTCGAGTATTAAAATAAATTATAAGTAATTGCAGAAGACTCCTTGAACAGACTCAAAAGGACTCCTTCGAATTTTACTCTTAATGCTACCTGAATTTTGTATTATAAATTTAAGGATCATGATTATGATATAAAGAATTTCTCAGCATATATGAAATGAACGAATTTGAGCATTGAACGTCTAACCTCATTTTTTAATTATTGTGATATGTAGAGTGGTGCAAAATCACAGATACCTCTCTTGATGCATTTCTATAGTTACGTTGATGTGAGATTATATATAGTTCAGCAGCAGCATGTTGGGAAAATTACTAATAACTCTTCTTTTATATCAAATTGTTGAAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGTTACCGGTTAGTGATACTCAGGTATTGTTTATCTACTTTATCATGTCGTAAGTATATTATTTGTAAAGATATATATCAAGATAGTTCGATTGCGTACACTTACATTTTGATTATGTTTGGTGAATACTATTCTAATACCTTTTTTTTTCCTAAAGCCTAACAAATAAAGATAATTAAGATGGGAACGTAATTCAAGTACAACATGGTTCCATACGTGACATATTTACACATATAGTGGAACCAAAAGAGCAATTTTTCCTAATATCATTTTCTAAATATCACGTGTGCCCGTGATTCTTTTTTATGGACATGAATTTTTTTTTTAATATGAGTGGAAGTAAGGTTCGATCTTTCTATCTGCTTTGATATCATATTGAATCGTGTGATTGTCTCTTTAAAAAATTAAGCAAGAGCATATTTTATTAATTAATTGTCTTTCTCGACGTTTTTCTCTTTCAACAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTCTCAAAGGTAAGATATTAGTGATGTAATTAAATGATTTTAGTTAGATTTACATAAGTTTTTAATAAGTGAAAATTAATAGACATATTCTTGGAGAGGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGGTATATCCAAATACTATAACTTAAATATATTGTAACGATTTAATTAATAGCATGTGTCACGTTCATCTATTCTTTAGTCACAATATATAGGGGCATGTCCTTAACAACGTGCCATGCCTCGATAGTCATTTTTGTCTTTTTGTGCGTATGAATTTAACTTTGACACAAATTTTTGTAGTAATAATAACTCATGCTTTAGCATCTTAGGAAGCAGTCATATGAAAAACAGAAGCATATATATATATTACATGAGTTAATTTAATTTAATATAAAATTTAATAAAATTGTGTCTCGCTATAAATAATTTTATTAAAAAATTATATAAATATATTATTTTTTTAACTGGCCGCAAAGTTATATAAATTGATAGAGAAAGAGGTTTTGGTGTAAGGTTCATTTTCCAACAATTAGTTTTATAATTTGTAAGTGCACACTTTATCAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGTACACTGCCTTAACATTACAAAATTAATTTATTTCATCAAAAGCATATCATAAAATTCTGACAAATAAATATATTAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGGTACATATCTATATATGTGTGTTAATTAATTAAGTTGATTTTGTATTTTTGTTTAATGAATAATTGTTTGTGATCATCAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGGTAAGTATCTTATTTTATATGACTTAGTTTGACTTGACATAAAGTTTAATAAAGAAAGAAAGACTTTTAAAACTTATAGTGTAAAATAAGTGAATAGATATATATGTGGTTGTACTAACACTACAACAAAAATAATTTTCAGCGGCATTAAATATTGACATTAATAATGAGTGCTAAAGACTTTATCGGTATTAGTTAAGTGTCATTAGGATCAATGTCGTTAAAGGCTTCACGGACATATACAAAGAGTGACAATTGCCGCTAATGATTATTTTTGTTGTAGTGAAAATGAGTATTTTAAAGTTAAATTGTTACATAATATAGAAATATGTCAGAAACAGGACAAATATACCACCGAACTATCATATATGTTATGGAGATATTCTCAGTCATACTTCTGCGACATTGGTACTCATGTCGTCCAAAAACTAGAACATATATATACCCTTTATATATTAACGAAGATACAAGTGTCATAATCTTATGCACCGATTCGATATTTATTAAATATCGAATCGACGGATAAAATTATGTCACGTGTCCCTATTAAGTCTTCTATTAGAGTAAAAAGCATATATTCTCTAGTTTTTGAACGAAAAAAGGTATTAATGTCTCAAAAGTATAACGAAAAGCATTTGCATACAATTTATGATAATTTGGGGCATATTAATTTATCATTCCCCCTTTTTTTGGCACTGATTAAAAAGAAAAAGAAAGTTATAAAAATTGGGATAGAGGGAATAATTGTTTCATAGGGAAAACTTAGAAGCTTCTCAGTATGTCAGTGAGAATGTGTTTCCTAATTAGTGAACTATGGTTTGGTGAAAAATAAAGAGAAAAAAATCAGTACAAATTTTCCACTGATTAGCAATGAGAAAAATATTTGTTTCTAGTAGTATGAGGAGAGGATAGTCCGCATAAATAATCCTTAAATTTGTGGATAAATAAACTATTTTCAATAGATTATCGTCTCAAAATAAAATAAAATGATTGCAAGAAAAGAATAATAGGTATGCTGGTAATATGTATAATACACTCAAATTTATTTGCTGTCCATGCAGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAAWild-type Solyc12g038510 coding sequence (SEQ ID NO: 7)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGTTACCGGTTAGTGATACTCAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTCTCAAAGACATATTCTTGGAGAGGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAA Mutant Solyc12g038510 gene allele j2^(TE) (SEQ ID NO: 8)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGGTATATATATATATACATATG

ATATGTTTTTCTTCTTTTTGTGTGTGCGTATGTGTTTACTTACTTTCATTAATTAACTCAACCATATATATACATCTCTCACCTCAATTATATATATGTTTGAGATCTGAATGTCTACGGACTCCATTTAGGTACATATCTTTGTTTAGATCATAAATCATCTATCTTCATTCCTAAGATCTACTAATATATATGTATAAGAAGATCCATCCATCTATTAGGTTTTTCAACAACATATACAGTGAAATCTTATATGTGGGCCCACGTATAGCCATATGAGAAAATAGTGTGCACGTAAACATTATCATTACTTAATTATAGGAATATACATCCATTAGGTTTATCAACAACAATAAAATCCTCTAAATGGAGTCTAGTCATAGGTCTAGCCGTTTGAAAATGTAAAATATATGCCGATCTTATCACTATGTCATAATAATAGATATGTTGTTATTGAAAGATTCTCAATCTTTTTTTTTCTTCAAGGTAGAGATTCTTAAGTGGATTCATGTTTTTTTTATCAAAAAAGAAAAAAACAAAAGTGTCCATTTGTTCATCTAATGGGTTTTCCATGTTACCAATTCACTACACTGTTGAGATTTGATTATCAGATGTGTCAAGTTTCGTTTGGTTCCCTAGAAGGGAGAAAAGGCTGCTTATGCAGGCAGGGTATTAAAGATGATATTAATATCTGCAGTAATCAGTAACAGAATATATAAACTTAATAATAAACTTGAAGGTACTTARTTATCCAGCAGATAATCTTCTGTCTCACCGTACACTTTTGTTATATCATAAGCATAAGAATTGTTTTATCAAATATTACCAAACAAAACTTAGTTTTGTTTGGTAATATTTTATAAAATATGTTACCGAAAGTTACTTCCTATAACATATTTTATAAAGAAAAAAATTAAAAACTCCATATACCTAAGAAATGTAACCCCCCCTCCATAACAACAATTTAACAAAAATAAAAACCTACTTTTTTTGAATTTGGTAAATTAGTTTTCTATCCTTTTTAGTAACTTCCTTTCTTATTTTCTTTTTATATTGGTAAAGTTTAATATTACACATTATTTTAACATGTTATAATTTTTTGTGATGCTTAATTATTTGATACATGTAATAAACCATATATTAGAGCTATAAATCAATGACAATGCATGTAGATACAACTCATTTATGATATATTTTGTTTATATATATAACCAATTAGATAATTTGTCTGCGCTTTGTGCAGTCATAAATAATAATTGCATTGAACTTGCAAATATTTTTTTTTAATATCCATACATTAAAAAAAAAGAAAGAGGAAAATTGGTTCCTAAAATATTAGCAATATTCAAACATTTATTTGATTATTAATCATTATCACATAACTTAAGAACGTCTAATGAATGAATTATTCACGAAATAATAAATCATTGGTTCTAAAAAGGAATTTCGTAATAAAATAAAAATTTAAGTTACCATATTCAAAAAAAGAAATTGTGCTTGAACATGAAAATAATTATAATTTTTGAACTTGTATAATGAATTTCTTCAATTCATAAGTGGGAAATTTCATATTTATGTAATAATAGATAATATGTAAGCTCTAATATAGTACTTTAGGTTATAGAATTTAATATAAAATATCAAAACATGAATTCTTGAAATTGAGTAGAGTAATTATTTTCTGCACAATGAATCGGAGACAATAACTTTGAAGAAATATAAACAATAGAGTTCAAAAGATGTAGTCAAAAACAACAATTAATATCATAAGAATAAATTAATGAGTGTAAAAATGCATACCACGATATGTAAAAACAGAATGGAATATAATAAAAAAAATCGAGTTCACTGAATACACAATGTTCCTTTAAGAAAATTATTCTCCTCCAATACCAACGAGATTACATCCTCTAAGGATGGAAATGATTTCATTCCCCAACTTATCCATATAAAAATAGTGGTGTTAGTATGTAACTCAATAGGAGTAAAATACACAAATATTTAATTTTGCGAAAGTAGAAGAAGAAGATCATATTTTTTTTTTAAAATGAGAGGATATATCACTATTTTTAAACAACAAAGGGTAGTGTTAACAAATTTTTATTGTGTCTTGTCTAAAAGGTTACAGCTATTTGAAAAAGTTACAACACTTCGAAAAGTGAACAACATTTCATAAAAGTCGTAACTTTTCATAAAGTCGTAACTCTTCATAAATGTCGCAACTCTTCATAAAAATTACAACTATTGATAAAAGTCACCACTCTTGATAAAGATCACCACTCTTCATTGAAGTTGCAACTTTTCATAAAAATCACATCTTTTAATAAAAAAGAAAGACTAGTTTTTGGAATAAATTAATTTAAAAGAAAATTTTTGTTTGTGGTGGGGCGCCAAGTAGGCAGGCGTAGGGTTCTTTTTATATAAATATATATGATATATGATTCAATATTTGATATATATATATATAGAGAGAGAGATGACAATATAAGACAATTGCAAAAAATAAAATAAAAAACTAATCGAGTAAGTAGGCAAAAAATTATTTATAAAATATATGTAGAATTTCTTTATCAGATATGACTGCCCAAATCTTATATTCAAACTAAAATGCAAGATCAATGGTGCTATATATAGGGTTTTACACAAAAATCAAGATCTAGTCTTGCAAATTTAAATAAAAAACAGTGGTTTACGATGAGATAATGTAGCTTTTGTAAACAATAAAACTAGAAAAATAAATGCAAAGGCATTTTAAAGGATATAATAATGAAGATCAAAGGCAGAGAAGGGAAGAGGCAGCAATATAATGAAGGTAACATCATGGTTCCATTCTAATATATATGCTATTTTTCTTTAGTAAATTTCAAAAATAATGATACATTTTCATATTTGATAAATATTTAATGATACTATCAACATTTTATCTATATTGAGTTCCATTTATTTGACCAAAACCTCACAAAGATGTGCTCTTCGATCTATTCAAAATTTATTCAATTTAAGGATAGCTTTAAAACATGACAAAGTTTTCTCATATATTTCTTAAATTTTATATCCAGTCTAAATACGTATATAAACTAAAATGAAGAGAATAATATGAAGCTTTATTTGATGACATTGTTGAAATAACCAAAAGCTATAAGTGATACAATAGTAAATTTACCATTGGTCAATTCAGAATTATTTAAAAGCTAAAAAAGTCATATAAGTTGGGGTTGCTCAATGTATAGTTTTTGGCTTGTTTTAAGCATTTTAAAACTTTTTTTAAGCGCTTTTTAACATTGCTAAACACTCAAAAAATGATAAATAGTATTTAAATTTGATATGATTAGCTTAAAAGTGAACTCATATACCTTCAAAGTAAAAATCCCCAATTCGAGCTTTCAAACCACTTGATTTTGTGGATGAAATTATACTGAAGTTGAATATATCACTATTTATAGGGGTTAGTGAACTAATACCTTTGATTATTTGGTAGAAATATGTATCTTAGATCACCCTAATGAGCTCCCACTTTTAAAATAGGAAAAACCTCATATGAAGTTCATCACTGTTCATTATATATCACTTTTATTCAAAAACGTTTACAAATGTTCATTGTGACTAAATACCCTTGAGTGTCGAGTTTTCACACCAATAAGGCCTAATTAATAGGTAAACAAAACTATGTCAATCTTCAAAACGCAAATCTAATTATATTTTTAACAAGATTAGAGGTATATATACATATTCTCTTATGTTAACTCTTATTCATTATTGAACAAACTAAGTAAGTGTACCCAAGGTCTCAAACAACAGTTGGTACATTCTTTGTATGTCTTCCTTTGTCTCTTAATAGTCGTCTCCTCCTGTCGATGATTCCTCCAAATACATTAATCAAAGGAAAATCTTTCGCCCTCAACTTGCAAACTTGTCTATCTAAAATTGTTAACAAAGTTTCTTCATTAGAGAAACTATGATTTCTTGAATGTAGCAATTTGATGTGCCATGACTATCATCTTGATCAACATGCTTCTTAACCATCAAAAGATCCTAAACTAGATGCATGTCATGTTAGGAGACATATTAAGCTTGTATATAACTACACCAACATGCTTTAGGATCTCATAAGATCCAAAATTTCTTATTTGGGAGATTTTCAATCCAACAACCATCATAATGAGCAACGTGATGTTATAACATCTCTCTCACACTGCCAGAACAGTCTTATACCTTGTCGGAGTGAAGGACATCCTTAACTAAGTAGATTCACTAAGCTATACTTAAAAAGCAATAAGGAATCATCTAAAATGTGTGACTCTTAACCCATATTGGCATACATGGTTTATGGGGGTTATTAATTGTCTGAACACTCCCCCATATAAATCAGTGATCAATATTAATCCCAATAATATACACTATTATGATTTGAGACTACACCCTGGAAGTGGCCGGCTCTCAAGAACCATTGCTGATCTCCAAGCCAAACCCTCATTCTGGTTGACTACAAGCTGAAGGCAAACTCAAGTATACAAAGCTTAAAACATAATAAAAATAATATACTCAACTCGCCACAAAATAGGCATTTAAGTCTTTAAAACATTTTTAAAAATAAATGAAACAAACTTCTCAAACTGTAATGTATATCTATGAAGCCTCTAAATGAAAAAAATGAAGGCAGATGAGACATACGGCATCCTAACAACTGATATAACTAAGAGTACAAGTGGAGCCCTTCGGATGTAAGGAGGCTCATCAAAGCTAATGTGAACTCCATGTGGTATCAATGAAGCACCTATTGATGACCGTGAATACATGTATCTGCATCATGAAACGATGCAGGCCAAAGGGCTTAGTACGTGAAATGTACGAGCATGTAAAGGGAATTCAAATACATAAACATAGGCTTGAACTTTGATATAAAGGAAACATACTTACCTATTTTTAACTCAAGAATAAAAAACATAGTTCAACTCAATGAAAAGACACTCAAGTCAGTGAAATAGGCCGCAACTCAATAATAAGATATTCGACTATGGGTAATCAACTCTGGGTACTCTATTCAATATAAAGTAAGAATACAAATGCATTATATGGAAAGACTTTAAAACGGTAGAAAACAACTCAATGTATTGAAAATTCAATAGTAAATTAGTTTGTATGTAAGGAACAATATAAACTTTGTTTGTATATGAAAATACAAAATAAACTTTGTGTATATAAAAGTACAAAATATCTCTGTGAAAGTTTCTCTAACCAACAACCATCACTATGAGCTTTCTGATAATACCACGTTTCGCCCATGATGTCAGAACTGTCCTATGATTTTCCAGTTCATAAGACCTACTCACTAAGTGGATCCACAAGTCTATGCTAAAAAATATTTAAGGAATCGTCTAAAAAGTATGACTCATTCTACCCACGTTGGCTACATGATTTATGGGGGTCGTAAGTTATCTAAACTCTCCTCCATATCGATGCGTAATGCTACTCACAAATATACTAGCTCACATGTTTAAAAATATAACTCGTTTTGTTTGAGATCATTACTCAAAATCCTTCTCTTAAAAGAGATGATACTCAAACTGCTCAAAACTCTTTTGGAAATCTCAAATTCGTCTCATCTTAAATGTAAAAATATTTACTCTTGGGAATACATAGTTATCATATATCATTTTAAAGAAAATGAACTCAACTCTGTTCTTTCTCAACTCAAGTGCTCAGTCTTAAACCAAATTAAAAAAAAGACTTCTCAAAATAAAGTTTATGTCGAATTATGGACGTGAACAATTCAATTCAAAGTTTTCGATAACCATAACTAAAACTAAATACTCGAGACTCAACATCTTAGAACTCAAGAACTTAAATGGTAATACTTCTTTCAAGAATGCTCGACTCAGAAGGTTAATGCAGAATAATGTGCATGAATTACTCAACTAAAGGACTCACTGATACTACTCAATCTCAAGATTGCTCGACTCGTAGGGTTAATGCAGAATTATGTGCATGAACTACTCAACTCAAAGACCTTCATAGGTAACATGTAGTAGCCCCATGATTTGGAATATAATCCCAAAATGATTAGGAACTCAATACTCAGGACTTAGAACTTGAAGATAATACTACTTCTCTCAAAGATACCCAACTGACGGAGTTCATGCAGAATTTATGGGCATGAACTACTCGACTCAAGAGTCTAAAACACAATATGACACTCATGTATATAACTCTTCTCATTCTAATACTTGTTTTCTCAAAACTCGGTTTAACTAAATAGTTGATCTCAAAGGATTCACAATTGAACTCAAAGACTTTCTTTGACTCCACTCTTAATTCTCTCTTAAATTTGTATTTGAATTATGAATTTAAGAGTTATGATTCATGATATGGGGAATCTCAATAACAATATAGAAATTTGATAATTAGGAATAGTACTTTTAAAAGAAAACATGAATTCAACTTAAAATCAACTTATCTAAAAAATATTCAAATATAGGGAAAGTATCCTAGACTACTGTGCTACTGATCTGAAAGTAGATGTAGGATGTGAGGATGAACTAGTCCAACACTATGATAGCCTTACATACCTGGAATAACGAGGTTCTTGGAAAATCTTCACTTGAAGAAGAACTTGATTAGAAGCCTTGAAACCTAGCTTGAAGGTAAACAATCAAGAAAACCTTTCTTAAGATTCTTGAATTAGTTTATGAAAATCTCTATGACCAAGCATTTTGATTTTCACTAGTGATTCATAATTGTATGGAGGAATTTGAATTGAAAAAGATGAAATGCTTGGAGAAAAGCTATCTTTGAAGAAGCTTGAAAAAGATTGGAAAGTCCTGTACTTTGATTTTCCCTTAGGATTTTGTCTTAGGGTTTGAGATAGAAAAGAATGATGGACTAAAAGATGAAAATCTAATTGTTTGGATCCTTTTTCAGCCAAGAAATCCGTTTAGGGTTTTCTTGGAGACAAACAAAATAAAAAAGACCATTTTTAATATTTTTCCGTCGGCTAATTCGTAATAACATTGTATCATGTTATTGAAAGAGTCATAACTTTTTACTCAAAAATTGGATTGATGCGAAATTAGTGGTGTTGGAAAGTAGATTCAAGTACCTCTAATTGGATAGGTTATTCCCTACATAAGTCTTTATATTCTAAAAGATATGGTTGTTTGCACTTGACCTAAGTAGAATTTTACATGAAAACTTAATAGAGAAGGAAACTTCAAGAACTCATCAAGAAATTTCAATTGCTCAATATTTATGGATAAATTTGTAGAAGAAACTCATGATTGACATGCGGGTGAATAAACCCAACACTATGGAAGCTTACATACCTCAAAGAACTAGGTTCTTGGCGAAATCTTGAATTTCTTCAACGAACGCTTGAAACTTTGAACTTTTTCTCTTCTTGAACTCTCAACTAAAACCCTAGGCGTATATTAGGATTATAAAAGTTAACATGATAGGATTAGACCTTTAAAAACTTTCTAAAATGAATTAAATCTGATTTAGCATGAAAAAGACCAAAATACCCCTTACTATTTTCGGATAACTTTTCTTAATTGGACTGCCTGACTTCAAAAAGGTATATCTCACTCATCCGACCTCAAAATTTAGCAAATTCAGTGGCGTTAGAAAGCTAATTTAAACACCTTTCATTTTCCATCTCATGGCACACATAACTCATTCTTTAAAGAGAGCTATGATCGTTCAAATTAACTCAAATCTTAGAAGAATTTAGGAATGTCTTGAACGAGCTACATCTAGTGACCTTAACACTTTGGAAAATTTTAAATTTCTTAGTAAAAACTTACTCACTATGAAGGATGGTTCAAGTCTTAGCTCAAAATTTTCCTAAGTTGCTATATATACTCATGCTCATATGTTTAAAACCAAAACCCTTCCTCGATTTGAATTAATTACCAAAAAGATTCTCTTAAAAAGATAATGCTCAAAACTCCCCCTAAACTCATTTGGAAATCTAGGTTTCCCTTGTTTTAAATATAAAAACATTTACTCTTGGAAATATTTAGTTCTCAGATATTCACTTGAAAAAAATTAAACTCGACTCTCATCATCTTCATACTCAAGTGCTCAAGTCCTAAAACAATTTATAACTAATTGTATAAGACTTCTCAAAATAGGGTTCATTCCGAATTATGGACGTGAACGACTCAATTCAAGGATTTCAATAACCATATATATAACTCAATAATAGGAACTCAACAACTCCAGAACTCAATGATACTACTCATCTCAAGAATGCTCGACTCACAGGGTCTTTGCGAAATTATTGGGCATGAACAACTCAACTCAAAGACCTTCATTTATACCATATGGTAGTCCCATAATAGGAATATAATCCCAAAAAAATTAGGAACTCAATACTCAAAAACTTAGAACTCGAAGATATTACTCATCTCAAAGATATTCAATTTATGGAATTCATGCTGAATTATGAGCATGAACGACTTGACTCAAGGATCTCAATAATAATGTAGACTCATGAATACACTCTTCTCATTCTCATACTCACATACTCGAGTATTAAAATAAATTATAAGTAATTGCAGAAGACTCCTTGAACAGACTCAAAAGGACTCCTTCGAATTTTACTCTTAATGCTACCTGAATTTTGTATTATAAATTTAAGGATCATGATTATGATATAAAGAATTTCTCAGCATATATGAAATGAACGAATTTGAGCATTGAACGTCTAACCTCATTTTTTAATTATTGTGATATGTAGAGTGGTGCAAAATCACAGATACCTCTCTTGATGCATTTCTATAGTTACGTTGATGTGAGATTATATATAGTTCAGCAGCAGCATGTTGGGAAAATTACTAATAACTCTTCTTTTATATCAAATTGTTGAAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGTTACCGGTTAGTGATACTCAGGTATTGTTTATCTACTTTATCATGTCGTAAGTATATTATTTGTAAAGATATATATCAAGATAGTTCGATTGCGTACACTTACATTTTGATTATGTTTGGTGAATACTATTCTAATACCTTTTTTTTTCCTAAAGCCTAACAAATAAAGATAATTAAGATGGGAACGTAATTCAAGTACAACATGGTTCCATACGTGACATATTTACACATATAGTGGAACCAAAAGAGCAATTTTTCCTAATATCATTTTCTAAATATCACGTGTGCCCGTGATTCTTTTTTATGGACATGAATTTTTTTTTTAATATGAGTGGAAGTAAGGTTCGATCTTTCTATCTGCTTTGATATCATATTGAATCGTGTGATTGTCTCTTTAAAAAATTAAGCAAGAGCATATTTTATTAATTAATTGTCTTTCTCGACGTTTTTCTCTTTCAACAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTCTCAAAGGTAAGATATTAGTGATGTAATTAAATGATTTTAGTTAGATTTACATAAGTTTTTAATAAGTGAAAATTAATAGACATATTCTTGGAGAGGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGGTATATCCAAATACTATAACTTAAATATATTGTAACGATTTAATTAATAGCATGTGTCACGTTCATCTATTCTTTAGTCACAATATATAGGGGCATGTCCTTAACAACGTGCCATGCCTCGATAGTCATTTTTGTCTTTTTGTGCGTATGAATTTAACTTTGACACAAATTTTTGTAGTAATAATAACTCATGCTTTAGCATCTTAGGAAGCAGTCATATGAAAAACAGAAGCATATATATATATTACATGAGTTAATTTAATTTAATATAAAATTTAATAAAATTGTGTCTCGCTATAAATAATTTTATTAAAAAATTATATAAATATATTATTTTTTTAACTGGCCGCAAAGTTATATAAATTGATAGAGAAAGAGGTTTTGGTGTAAGGTTCATTTTCCAACAATTAGTTTTATAATTTGTAAGTGCACACTTTATCAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGTACACTGCCTTAACATTACAAAATTAATTTATTTCATCAAAAGCATATCATAAAATTCTGACAAATAAATATATTAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGGTACATATCTATATATGTGTGTTAATTAATTAAGTTGATTTTGTATTTTTGTTTAATGAATAATTGTTTGTGATCATCAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGGTAAGTATCTTATTTTATATGACTTAGTTTGACTTGACATAAAGTTTAATAAAGAAAGAAAGACTTTTAAAACTTATAGTGTAAAATAAGTGAATAGATATATATGTGGTTGTACTAACACTACAACAAAAATAATTTTCAGCGGCATTAAATATTGACATTAATAATGAGTGCTAAAGACTTTATCGGTATTAGTTAAGTGTCATTAGGATCAATGTCGTTAAAGGCTTCACGGACATATACAAAGAGTGACAATTGCCGCTAATGATTATTTTTGTTGTAGTGAAAATGAGTATTTTAAAGTTAAATTGTTACATAATATAGAAATATGTCAGAAACAGGACAAATATACCACCGAACTATCATATATGTTATGGAGATATTCTCAGTCATACTTCTGCGACATTGGTACTCATGTCGTCCAAAAACTAGAACATATATATACCCTTTATATATTAACGAAGATACAAGTGTCATAATCTTATGCACCGATTCGATATTTATTAAATATCGAATCGACGGATAAAATTATGTCACGTGTCCCTATTAAGTCTTCTATTAGAGTAAAAAGCATATATTCTCTAGTTTTTGAACGAAAAAAGGTATTAATGTCTCAAAAGTATAACGAAAAGCATTTGCATACAATTTATGATAATTTGGGGCATATTAATTTATCATTCCCCCTTTTTTTGGCACTGATTAAAAAGAAAAAGAAAGTTATAAAAATTGGGATAGAGGGAATAATTGTTTCATAGGGAAAACTTAGAAGCTTCTCAGTATGTCAGTGAGAATGTGTTTCCTAATTAGTGAACTATGGTTTGGTGAAAAATAAAGAGAAAAAAATCAGTACAAATTTTCCACTGATTAGCAATGAGAAAAATATTTGTTTCTAGTAGTATGAGGAGAGGATAGTCCGCATAAATAATCCTTAAATTTGTGGATAAATAAACTATTTTCAATAGATTATCGTCTCAAAATAAAATAAAATGATTGCAAGAAAAGAATAATAGGTATGCTGGTAATATGTATAATACACTCAAATTTATTTGCTGTCCATGCAGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAAMutant Solyc12g038510 gene allele j2^(stop) (SEQ ID NO: 9)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGGTATATATATATATACATATGTTTTTCTTCTTTTTGTGTGTGCGTATGTGTTTACTTACTTTCATTAATTAACTCAACCATATATATACATCTCTCACCTCAATTATATATATGTTTGAGATCTGAATGTCTACGGACTCCATTTAGGTACATATCTTTGTTTAGATCATAAATCATCTATCTTCATTCCTAAGATCTACTAATATATATGTATAAGAAGATCCATCCATCTATTAGGTTTTTCAACAACATATACAGTGAAATCTTATATGTGGGCCCACGTATAGCCATATGAGAAAATAGTGTGCACGTAAACATTATCATTACTTAATTATAGGAATATACATCCATTAGGTTTATCAACAACAATAAAATCCTCTAAATGGAGTCTAGTCATAGGTCTAGCCGTTTGAAAATGTAAAATATATGCCGATCTTATCACTATGTCATAATAATAGATATGTTGTTATTGAAAGATTCTCAATCTTTTTTTTTCTTCAAGGTAGAGATTCTTAAGTGGATTCATGTTTTTTTTATCAAAAAAGAAAAAAACAAAAGTGTCCATTTGTTCATCTAATGGGTTTTCCATGTTACCAATTCACTACACTGTTGAGATTTGATTATCAGATGTGTCAAGTTTCGTTTGGTTCCCTAGAAGGGAGAAAAGGCTGCTTATGCAGGCAGGGTATTAAAGATGATATTAATATCTGCAGTAATCAGTAACAGAATATATAAACTTAATAATAAACTTGAAGGTACTTAATTATCCAGCAGATAATCTTCTGTCTCACCGTACACTTTTGTTATATCATAAGCATAAGAATTGTTTTATCAAATATTACCAAACAAAACTTAGTTTTGTTTGGTAATATTTTATAAAATATGTTACCGAAAGTTACTTCCTATAACATATTTTATAAAGAAAAAAATTAAAAACTCCATATACCTAAGAAATGTAACCCCCCCTCCATAACAACAATTTAACAAAAATAAAAACCTACTTTTTTTGAATTTGGTAAATTAGTTTTCTATCCTTTTTAGTAACTTCCTTTCTTATTTTCTTTTTATATTGGTAAAGTTTAATATTACACATTATTTTAACATGTTATAATTTTTTGTGATGCTTAATTATTTGATACATGTAATAAACCATATATTAGAGCTATAAATCAATGACAATGCATGTAGATACAACTCATTTATGATATATTTTGTTTATATATATAACCAATTAGATAATTTGTCTGCGCTTTGTGCAGTCATAAATAATAATTGCATTGAACTTGCAAATATTTTTTTTTAATATCCATACATTAAAAAAAAAGAAAGAGGAAAATTGGTTCCTAAAATATTAGCAATATTCAAACATTTATTTGATTATTAATCATTATCACATAACTTAAGAACGTCTAATGAATGAATTATTCACGAAATAATAAATCATTGGTTCTAAAAAGGAATTTCGTAATAAAATAAAAATTTAAGTTACCATATTCAAAAAAAGAAATTGTGCTTGAACATGAAAATAATTATAATTTTTGAACTTGTATAATGAATTTCTTCAATTCATAAGTGGGAAATTTCATATTTATGTAATAATAGATAATATGTAAGCTCTAATATAGTACTTTAGGTTATAGAATTTAATATAAAATATCAAAACATGAATTCTTGAAATTGAGTAGAGTAATTATTTTCTGCACAATGAATCGGAGACAATAACTTTGAAGAAATATAAACAATAGAGTTCAAAAGATGTAGTCAAAAACAACAATTAATATCATAAGAATAAATTAATGAGTGTAAAAATGCATACCACGATATGTAAAAACAGAATGGAATATAATAAAAAAAATCGAGTTCACTGAATACACAATGTTCCTTTAAGAAAATTATTCTCCTCCAATACCAACGAGATTACATCCTCTAAGGATGGAAATGATTTCATTCCCCAACTTATCCATATAAAAATAGTGGTGTTAGTATGTAACTCAATAGGAGTAAAATACACAAATATTTAATTTTGCGAAAGTAGAAGAAGAAGATCATATTTTTTTTTTAAAATGAGAGGATATATCACTATTTTTAAACAACAAAGGGTAGTGTTAACAAATTTTTATTGTGTCTTGTCTAAAAGGTTACAGCTATTTGAAAAAGTTACAACACTTCGAAAAGTGAACAACATTTCATAAAAGTCGTAACTTTTCATAAAGTCGTAACTCTTCATAAATGTCGCAACTCTTCATAAAAATTACAACTATTGATAAAAGTCACCACTCTTGATAAAGATCACCACTCTTCATTGAAGTTGCAACTTTTCATAAAAATCACATCTTTTAATAAAAAAGAAAGACTAGTTTTTGGAATAAATTAATTTAAAAGAAAATTTTTGTTTGTGGTGGGGCGCCAAGTAGGCAGGCGTAGGGTTCTTTTTATATAAATATATATGATATATGATTCAATATTTGATATATATATATATAGAGAGAGAGATGACAATATAAGACAATTGCAAAAAATAAAATAAAAAACTAATCGAGTAAGTAGGCAAAAAATTATTTATAAAATATATGTAGAATTTCTTTATCAGATATGACTGCCCAAATCTTATATTCAAACTAAAATGCAAGATCAATGGTGCTATATATAGGGTTTTACACAAAAATCAAGATCTAGTCTTGCAAATTTAAATAAAAAACAGTGGTTTACGATGAGATAATGTAGCTTTTGTAAACAATAAAACTAGAAAAATAAATGCAAAGGCATTTTAAAGGATATAATAATGAAGATCAAAGGCAGAGAAGGGAAGAGGCAGCAATATAATGAAGGTAACATCATGGTTCCATTCTAATATATATGCTATTTTTCTTTAGTAAATTTCAAAAATAATGATACATTTTCATATTTGATAAATATTTAATGATACTATCAACATTTTATCTATATTGAGTTCCATTTATTTGACCAAAACCTCACAAAGATGTGCTCTTCGATCTATTCAAAATTTATTCAATTTAAGGATAGCTTTAAAACATGACAAAGTTTTCTCATATATTTCTTAAATTTTATATCCAGTCTAAATACGTATATAAACTAAAATGAAGAGAATAATATGAAGCTTTATTTGATGACATTGTTGAAATAACCAAAAGCTATAAGTGATACAATAGTAAATTTACCATTGGTCAATTCAGAATTATTTAAAAGCTAAAAAAGTCATATAAGTTGGGGTTGCTCAATGTATAGTTTTTGGCTTGTTTTAAGCATTTTAAAACTTTTTTTAAGCGCTTTTTAACATTGCTAAACACTCAAAAAATGATAAATAGTATTTAAATTTGATATGATTAGCTTAAAAGTGAACTCATATACCTTCAAAGTAAAAATCCCCAATTCGAGCTTTCAAACCACTTGATTTTGTGGATGAAATTATACTGAAGTTGAATATATCACTATTTATAGGGGTTAGTGAACTAATACCTTTGATTATTTGGTAGAAATATGTATCTTAGATCACCCTAATGAGCTCCCACTTTTAAAATAGGAAAAACCTCATATGAAGTTCATCACTGTTCATTATATATCACTTTTATTCAAAAACGTTTACAAATGTTCATTGTGACTAAATACCCTTGAGTGTCGAGTTTTCACACCAATAAGGCCTAATTAATAGGTAAACAAAACTATGTCAATCTTCAAAACGCAAATCTAATTATATTTTTAACAAGATTAGAGGTATATATACATATTCTCTTATGTTAACTCTTATTCATTATTGAACAAACTAAGTAAGTGTACCCAAGGTCTCAAACAACAGTTGGTACATTCTTTGTATGTCTTCCTTTGTCTCTTAATAGTCGTCTCCTCCTGTCGATGATTCCTCCAAATACATTAATCAAAGGAAAATCTTTCGCCCTCAACTTGCAAACTTGTCTATCTAAAATTGTTAACAAAGTTTCTTCATTAGAGAAACTATGATTTCTTGAATGTAGCAATTTGATGTGCCATGACTATCATCTTGATCAACATGCTTCTTAACCATCAAAAGATCCTAAACTAGATGCATGTCATGTTAGGAGACATATTAAGCTTGTATATAACTACACCAACATGCTTTAGGATCTCATAAGATCCAAAATTTCTTATTTGGGAGATTTTCAATCCAACAACCATCATAATGAGCAACGTGATGTTATAACATCTCTCTCACACTGCCAGAACAGTCTTATACCTTGTCGGAGTGAAGGACATCCTTAACTAAGTAGATTCACTAAGCTATACTTAAAAAGCAATAAGGAATCATCTAAAATGTGTGACTCTTAACCCATATTGGCATACATGGTTTATGGGGGTTATTAATTGTCTGAACACTCCCCCATATAAATCAGTGATCAATATTAATCCCAATAATATACACTATTATGATTTGAGACTACACCCTGGAAGTGGCCGGCTCTCAAGAACCATTGCTGATCTCCAAGCCAAACCCTCATTCTGGTTGACTACAAGCTGAAGGCAAACTCAAGTATACAAAGCTTAAAACATAATAAAAATAATATACTCAACTCGCCACAAAATAGGCATTTAAGTCTTTAAAACATTTTTAAAAATAAATGAAACAAACTTCTCAAACTGTAATGTATATCTATGAAGCCTCTAAATGAAAAAAATGAAGGCAGATGAGACATACGGCATCCTAACAACTGATATAACTAAGAGTACAAGTGGAGCCCTTCGGATGTAAGGAGGCTCATCAAAGCTAATGTGAACTCCATGTGGTATCAATGAAGCACCTATTGATGACCGTGAATACATGTATCTGCATCATGAAACGATGCAGGCCAAAGGGCTTAGTACGTGAAATGTACGAGCATGTAAAGGGAATTCAAATACATAAACATAGGCTTGAACTTTGATATAAAGGAAACATACTTACCTATTTTTAACTCAAGAATAAAAAACATAGTTCAACTCAATGAAAAGACACTCAAGTCAGTGAAATAGGCCGCAACTCAATAATAAGATATTCGACTATGGGTAATCAACTCTGGGTACTCTATTCAATATAAAGTAAGAATACAAATGCATTATATGGAAAGACTTTAAAACGGTAGAAAACAACTCAATGTATTGAAAATTCAATAGTAAATTAGTTTGTATGTAAGGAACAATATAAACTTTGTTTGTATATGAAAATACAAAATAAACTTTGTGTATATAAAAGTACAAAATATCTCTGTGAAAGTTTCTCTAACCAACAACCATCACTATGAGCTTTCTGATAATACCACGTTTCGCCCATGATGTCAGAACTGTCCTATGATTTTCCAGTTCATAAGACCTACTCACTAAGTGGATCCACAAGTCTATGCTAAAAAATATTTAAGGAATCGTCTAAAAAGTATGACTCATTCTACCCACGTTGGCTACATGATTTATGGGGGTCGTAAGTTATCTAAACTCTCCTCCATATCGATGCGTAATGCTACTCACAAATATACTAGCTCACATGTTTAAAAATATAACTCGTTTTGTTTGAGATCATTACTCAAAATCCTTCTCTTAAAAGAGATGATACTCAAACTGCTCAAAACTCTTTTGGAAATCTCAAATTCGTCTCATCTTAAATGTAAAAATATTTACTCTTGGGAATACATAGTTATCATATATCATTTTAAAGAAAATGAACTCAACTCTGTTCTTTCTCAACTCAAGTGCTCAGTCTTAAACCAAATTAAAAAAAAGACTTCTCAAAATAAAGTTTATGTCGAATTATGGACGTGAACAATTCAATTCAAAGTTTTCGATAACCATAACTAAAACTAAATACTCGAGACTCAACATCTTAGAACTCAAGAACTTAAATGGTAATACTTCTTTCAAGAATGCTCGACTCAGAAGGTTAATGCAGAATAATGTGCATGAATTACTCAACTAAAGGACTCACTGATACTACTCAATCTCAAGATTGCTCGACTCGTAGGGTTAATGCAGAATTATGTGCATGAACTACTCAACTCAAAGACCTTCATAGGTAACATGTAGTAGCCCCATGATTTGGAATATAATCCCAAAATGATTAGGAACTCAATACTCAGGACTTAGAACTTGAAGATAATACTACTTCTCTCAAAGATACCCAACTGACGGAGTTCATGCAGAATTTATGGGCATGAACTACTCGACTCAAGAGTCTAAAACACAATATGACACTCATGTATATAACTCTTCTCATTCTAATACTTGTTTTCTCAAAACTCGGTTTAACTAAATAGTTGATCTCAAAGGATTCACAATTGAACTCAAAGACTTTCTTTGACTCCACTCTTAATTCTCTCTTAAATTTGTATTTGAATTATGAATTTAAGAGTTATGATTCATGATATGGGGAATCTCAATAACAATATAGAAATTTGATAATTAGGAATAGTACTTTTAAAAGAAAACATGAATTCAACTTAAAATCAACTTATCTAAAAAATATTCAAATATAGGGAAAGTATCCTAGACTACTGTGCTACTGATCTGAAAGTAGATGTAGGATGTGAGGATGAACTAGTCCAACACTATGATAGCCTTACATACCTGGAATAACGAGGTTCTTGGAAAATCTTCACTTGAAGAAGAACTTGATTAGAAGCCTTGAAACCTAGCTTGAAGGTAAACAATCAAGAAAACCTTTCTTAAGATTCTTGAATTAGTTTATGAAAATCTCTATGACCAAGCATTTTGATTTTCACTAGTGATTCATAATTGTATGGAGGAATTTGAATTGAAAAAGATGAAATGCTTGGAGAAAAGCTATCTTTGAAGAAGCTTGAAAAAGATTGGAAAGTCCTGTACTTTGATTTTCCCTTAGGATTTTGTCTTAGGGTTTGAGATAGAAAAGAATGATGGACTAAAAGATGAAAATCTAATTGTTTGGATCCTTTTTCAGCCAAGAAATCCGTTTAGGGTTTTCTTGGAGACAAACAAAATAAAAAAGACCATTTTTAATATTTTTCCGTCGGCTAATTCGTAATAACATTGTATCATGTTATTGAAAGAGTCATAACTTTTTACTCAAAAATTGGATTGATGCGAAATTAGTGGTGTTGGAAAGTAGATTCAAGTACCTCTAATTGGATAGGTTATTCCCTACATAAGTCTTTATATTCTAAAAGATATGGTTGTTTGCACTTGACCTAAGTAGAATTTTACATGAAAACTTAATAGAGAAGGAAACTTCAAGAACTCATCAAGAAATTTCAATTGCTCAATATTTATGGATAAATTTGTAGAAGAAACTCATGATTGACATGCGGGTGAATAAACCCAACACTATGGAAGCTTACATACCTCAAAGAACTAGGTTCTTGGCGAAATCTTGAATTTCTTCAACGAACGCTTGAAACTTTGAACTTTTTCTCTTCTTGAACTCTCAACTAAAACCCTAGGCGTATATTAGGATTATAAAAGTTAACATGATAGGATTAGACCTTTAAAAACTTTCTAAAATGAATTAAATCTGATTTAGCATGAAAAAGACCAAAATACCCCTTACTATTTTCGGATAACTTTTCTTAATTGGACTGCCTGACTTCAAAAAGGTATATCTCACTCATCCGACCTCAAAATTTAGCAAATTCAGTGGCGTTAGAAAGCTAATTTAAACACCTTTCATTTTCCATCTCATGGCACACATAACTCATTCTTTAAAGAGAGCTATGATCGTTCAAATTAACTCAAATCTTAGAAGAATTTAGGAATGTCTTGAACGAGCTACATCTAGTGACCTTAACACTTTGGAAAATTTTAAATTTCTTAGTAAAAACTTACTCACTATGAAGGATGGTTCAAGTCTTAGCTCAAAATTTTCCTAAGTTGCTATATATACTCATGCTCATATGTTTAAAACCAAAACCCTTCCTCGATTTGAATTAATTACCAAAAAGATTCTCTTAAAAAGATAATGCTCAAAACTCCCCCTAAACTCATTTGGAAATCTAGGTTTCCCTTGTTTTAAATATAAAAACATTTACTCTTGGAAATATTTAGTTCTCAGATATTCACTTGAAAAAAATTAAACTCGACTCTCATCATCTTCATACTCAAGTGCTCAAGTCCTAAAACAATTTATAACTAATTGTATAAGACTTCTCAAAATAGGGTTCATTCCGAATTATGGACGTGAACGACTCAATTCAAGGATTTCAATAACCATATATATAACTCAATAATAGGAACTCAACAACTCCAGAACTCAATGATACTACTCATCTCAAGAATGCTCGACTCACAGGGTCTTTGCGAAATTATTGGGCATGAACAACTCAACTCAAAGACCTTCATTTATACCATATGGTAGTCCCATAATAGGAATATAATCCCAAAAAAATTAGGAACTCAATACTCAAAAACTTAGAACTCGAAGATATTACTCATCTCAAAGATATTCAATTTATGGAATTCATGCTGAATTATGAGCATGAACGACTTGACTCAAGGATCTCAATAATAATGTAGACTCATGAATACACTCTTCTCATTCTCATACTCACATACTCGAGTATTAAAATAAATTATAAGTAATTGCAGAAGACTCCTTGAACAGACTCAAAAGGACTCCTTCGAATTTTACTCTTAATGCTACCTGAATTTTGTATTATAAATTTAAGGATCATGATTATGATATAAAGAATTTCTCAGCATATATGAAATGAACGAATTTGAGCATTGAACGTCTAACCTCATTTTTTAATTATTGTGATATGTAGAGTGGTGCAAAATCACAGATACCTCTCTTGATGCATTTCTATAGTTACGTTGATGTGAGATTATATATAGTTCAGCAGCAGCATGTTGGGAAAATTACTAATAACTCTTCTTTTATATCAAATTGTTGAAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGT

ACCGGTTAGTGATACTCAGGTATTGTTTATCTACTTTATCATGTCGTAAGTATATTATTTGTAAAGATATATATCAAGATAGTTCGATTGCGTACACTTACATTTTGATTATGTTTGGTGAATACTATTCTAATACCTTTTTTTTTCCTAAAGCCTAACAAATAAAGATAATTAAGATGGGAACGTAATTCAAGTACAACATGGTTCCATACGTGACATATTTACACATATAGTGGAACCAAAAGAGCAATTTTTCCTAATATCATTTTCTAAATATCACGTGTGCCCGTGATTCTTTTTTATGGACATGAATTTTTTTTTTAATATGAGTGGAAGTAAGGTTCGATCTTTCTATCTGCTTTGATATCATATTGAATCGTGTGATTGTCTCTTTAAAAAATTAAGCAAGAGCATATTTTATTAATTAATTGTCTTTCTCGACGTTTTTCTCTTTCAACAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTCTCAAAGGTAAGATATTAGTGATGTAATTAAATGATTTTAGTTAGATTTACATAAGTTTTTAATAAGTGAAAATTAATAGACATATTCTTGGAGAGGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGGTATATCCAAATACTATAACTTAAATATATTGTAACGATTTAATTAATAGCATGTGTCACGTTCATCTATTCTTTAGTCACAATATATAGGGGCATGTCCTTAACAACGTGCCATGCCTCGATAGTCATTTTTGTCTTTTTGTGCGTATGAATTTAACTTTGACACAAATTTTTGTAGTAATAATAACTCATGCTTTAGCATCTTAGGAAGCAGTCATATGAAAAACAGAAGCATATATATATATTACATGAGTTAATTTAATTTAATATAAAATTTAATAAAATTGTGTCTCGCTATAAATAATTTTATTAAAAAATTATATAAATATATTATTTTTTTAACTGGCCGCAAAGTTATATAAATTGATAGAGAAAGAGGTTTTGGTGTAAGGTTCATTTTCCAACAATTAGTTTTATAATTTGTAAGTGCACACTTTATCAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGTACACTGCCTTAACATTACAAAATTAATTTATTTCATCAAAAGCATATCATAAAATTCTGACAAATAAATATATTAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGGTACATATCTATATATGTGTGTTAATTAATTAAGTTGATTTTGTATTTTTGTTTAATGAATAATTGTTTGTGATCATCAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGGTAAGTATCTTATTTTATATGACTTAGTTTGACTTGACATAAAGTTTAATAAAGAAAGAAAGACTTTTAAAACTTATAGTGTAAAATAAGTGAATAGATATATATGTGGTTGTACTAACACTACAACAAAAATAATTTTCAGCGGCATTAAATATTGACATTAATAATGAGTGCTAAAGACTTTATCGGTATTAGTTAAGTGTCATTAGGATCAATGTCGTTAAAGGCTTCACGGACATATACAAAGAGTGACAATTGCCGCTAATGATTATTTTTGTTGTAGTGAAAATGAGTATTTTAAACTTAAATTGTTACATAATATAGAAATATGTCAGAAACAGGACAAATATACCACCGAACTATCATATATGTTATGGAGATATTCTCAGTCATACTTCTGCGACATTGGTACTCATGTCGTCCAAAAACTAGAACATATATATACCCTTTATATATTAACGAAGATACAAGTGTCATAATCTTATGCACCGATTCGATATTTATTAAATATCGAATCGACGGATAAAATTATGTCACGTGTCCCTATTAAGTCTTCTATTAGAGTAAAAAGCATATATTCTCTAGTTTTTGAACGAAAAAAGGTATTAATGTCTCAAAAGTATAACGAAAAGCATTTGCATACAATTTATGATAATTTGGGGCATATTAATTTATCATTCCCCCTTTTTTTGGCACTGATTAAAAAGAAAAAGAAAGTTATAAAAATTGGGATAGAGGGAATAATTGTTTCATAGGGAAAACTTAGAAGCTTCTCAGTATGTCAGTGAGAATGTGTTTCCTAATTAGTGAACTATGGTTTGGTGAAAAATAAAGAGAAAAAAATCAGTACAAATTTTCCACTGATTAGCAATGAGAAAAATATTTGTTTCTAGTAGTATGAGGAGAGGATAGTCCGCATAAATAATCCTTAAATTTGTGGATAAATAAACTATTTTCAATAGATTATCGTCTCAAAATAAAATAAAATGATTGCAAGAAAAGAATAATAGGTATGCTGGTAATATGTATAATACACTCAAATTTATTTGCTGTCCATGCAGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAAMutant Solyc12g038510 gene allele j2^(CR) >allele-1 (SEQ ID NO: 10)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGGTATATATATATATACATATGTTTTTCTTCTTTTTGTGTGTGCGTATGTGTTTACTTACTTTCATTAATTAACTCAACCATATATATACATCTCTCACCTCAATTATATATATGTTTGAGATCTGAATGTCTACGGACTCCATTTAGGTACATATCTTTGTTTAGATCATAAATCATCTATCTTCATTCCTAAGATCTACTAATATATATGTATAAGAAGATCCATCCATCTATTAGGTTTTTCAACAACATATACAGTGAAATCTTATATGTGGGCCCACGTATAGCCATATGAGAAAATAGTGTGCACGTAAACATTATCATTACTTAATTATAGGAATATACATCCATTAGGTTTATCAACAACAATAAAATCCTCTAAATGGAGTCTAGTCATAGGTCTAGCCGTTTGAAAATGTAAAATATATGCCGATCTTATCACTATGTCATAATAATAGATATGTTGTTATTGAAAGATTCTCAATCTTTTTTTTTCTTCAAGGTAGAGATTCTTAAGTGGATTCATGTTTTTTTTATCAAAAAAGAAAAAAACAAAAGTGTCCATTTGTTCATCTAATGGGTTTTCCATGTTACCAATTCACTACACTGTTGAGATTTGATTATCAGATGTGTCAAGTTTCGTTTGGTTCCCTAGAAGGGAGAAAAGGCTGCTTATGCAGGCAGGGTATTAAAGATGATATTAATATCTGCAGTAATCAGTAACAGAATATATAAACTTAATAATAAACTTGAAGGTACTTAATTATCCAGCAGATAATCTTCTGTCTCACCGTACACTTTTGTTATATCATAAGCATAAGAATTGTTTTATCAAATATTACCAAACAAAACTTAGTTTTGTTTGGTAATATTTTATAAAATATGTTACCGAAAGTTACTTCCTATAACATATTTTATAAAGAAAAAAATTAAAAACTCCATATACCTAAGAAATGTAACCCCCCCTCCATAACAACAATTTAACAAAAATAAAAACCTACTTTTTTTGAATTTGGTAAATTAGTTTTCTATCCTTTTTAGTAACTTCCTTTCTTATTTTCTTTTTATATTGGTAAAGTTTAATATTACACATTATTTTAACATGTTATAATTTTTTGTGATGCTTAATTATTTGATACATGTAATAAACCATATATTAGAGCTATAAATCAATGACAATGCATGTAGATACAACTCATTTATGATATATTTTGTTTATATATATAACCAATTAGATAATTTGTCTGCGCTTTGTGCAGTCATAAATAATAATTGCATTGAACTTGCAAATATTTTTTTTTAATATCCATACATTAAAAAAAAAGAAAGAGGAAAATTGGTTCCTAAAATATTAGCAATATTCAAACATTTATTTGATTATTAATCATTATCACATAACTTAAGAACGTCTAATGAATGAATTATTCACGAAATAATAAATCATTGGTTCTAAAAAGGAATTTCGTAATAAAATAAAAATTTAAGTTACCATATTCAAAAAAAGAAATTGTGCTTGAACATGAAAATAATTATAATTTTTGAACTTGTATAATGAATTTCTTCAATTCATAAGTGGGAAATTTCATATTTATGTAATAATAGATAATATGTAAGCTCTAATATAGTACTTTAGGTTATAGAATTTAATATAAAATATCAAAACATGAATTCTTGAAATTGAGTAGAGTAATTATTTTCTGCACAATGAATCGGAGACAATAACTTTGAAGAAATATAAACAATAGAGTTCAAAAGATGTAGTCAAAAACAACAATTAATATCATAAGAATAAATTAATGAGTGTAAAAATGCATACCACGATATGTAAAAACAGAATGGAATATAATAAAAAAAATCGAGTTCACTGAATACACAATGTTCCTTTAAGAAAATTATTCTCCTCCAATACCAACGAGATTACATCCTCTAAGGATGGAAATGATTTCATTCCCCAACTTATCCATATAAAAATAGTGGTGTTAGTATGTAACTCAATAGGAGTAAAATACACAAATATTTAATTTTGCGAAAGTAGAAGAAGAAGATCATATTTTTTTTTTAAAATGAGAGGATATATCACTATTTTTAAACAACAAAGGGTAGTGTTAACAAATTTTTATTGTGTCTTGTCTAAAAGGTTACAGCTATTTGAAAAAGTTACAACACTTCGAAAAGTGAACAACATTTCATAAAAGTCGTAACTTTTCATAAAGTCGTAACTCTTCATAAATGTCGCAACTCTTCATAAAAATTACAACTATTGATAAAAGTCACCACTCTTGATAAAGATCACCACTCTTCATTGAAGTTGCAACTTTTCATAAAAATCACATCTTTTAATAAAAAAGAAAGACTAGTTTTTGGAATAAATTAATTTAAAAGAAAATTTTTGTTTGTGGTGGGGCGCCAAGTAGGCAGGCGTAGGGTTCTTTTTATATAAATATATATGATATATGATTCAATATTTGATATATATATATATAGAGAGAGAGATGACAATATAAGACAATTGCAAAAAATAAAATAAAAAACTAATCGAGTAAGTAGGCAAAAAATTATTTATAAAATATATGTAGAATTTCTTTATCAGATATGACTGCCCAAATCTTATATTCAAACTAAAATGCAAGATCAATGGTGCTATATATAGGGTTTTACACAAAAATCAAGATCTAGTCTTGCAAATTTAAATAAAAAACAGTGGTTTACGATGAGATAATGTAGCTTTTGTAAACAATAAAACTAGAAAAATAAATGCAAAGGCATTTTAAAGGATATAATAATGAAGATCAAAGGCAGAGAAGGGAAGAGGCAGCAATATAATGAAGGTAACATCATGGTTCCATTCTAATATATATGCTATTTTTCTTTAGTAAATTTCAAAAATAATGATACATTTTCATATTTGATAAATATTTAATGATACTATCAACATTTTATCTATATTGAGTTCCATTTATTTGACCAAAACCTCACAAAGATGTGCTCTTCGATCTATTCAAPATTTATTCAATTTAAGGATAGCTTTAAAACATGACAAAGTTTTCTCATATATTTCTTAAATTTTATATCCAGTCTAAATACGTATATAAACTAAAATGAAGAGAATAATATGAAGCTTTATTTGATGACATTGTTGAAATAACCAAAAGCTATAAGTGATACAATAGTAAATTTACCATTGGTCAATTCAGAATTATTTAAAAGCTAAAAAAGTCATATAAGTTGGGGTTGCTCAATGTATAGTTTTTGGCTTGTTTTAAGCATTTTAAAACTTTTTTTAAGCGCTTTTTAACATTGCTAAACACTCAAAAAATGATAAATAGTATTTAAATTTGATATGATTAGCTTAAAAGTGAACTCATATACCTTCAAAGTAAAAATCCCCAATTCGAGCTTTCAAACCACTTGATTTTGTGGATGAAATTATACTGAAGTTGAATATATCACTATTTATAGGGGTTAGTGAACTAATACCTTTGATTATTTGGTAGAAATATGTATCTTAGATCACCCTAATGAGCTCCCACTTTTAAAATAGGAAAAACCTCATATGAAGTTCATCACTGTTCATTATATATCACTTTTATTCAAAAACGTTTACAAATGTTCATTGTGACTAAATACCCTTGAGTGTCGAGTTTTCACACCAATAAGGCCTAATTAATAGGTAAACAAAACTATGTCAATCTTCAAAACGCAAATCTAATTATATTTTTAACAAGATTAGAGGTATATATACATATTCTCTTATGTTAACTCTTATTCATTATTGAACAAACTAAGTAAGTGTACCCAAGGTCTCAAACAACAGTTGGTACATTCTTTGTATGTCTTCCTTTGTCTCTTAATAGTCGTCTCCTCCTGTCGATGATTCCTCCAAATACATTAATCAAAGGAAAATCTTTCGCCCTCAACTTGCAAACTTGTCTATCTAAAATTGTTAACAAAGTTTCTTCATTAGAGAAACTATGATTTCTTGAATGTAGCAATTTGATGTGCCATGACTATCATCTTGATCAACATGCTTCTTAACCATCAAAAGATCCTAAACTAGATGCATGTCATGTTAGGAGACATATTAAGCTTGTATATAACTACACCAACATGCTTTAGGATCTCATAAGATCCAAAATTTCTTATTTGGGAGATTTTCAATCCAACAACCATCATAATGAGCAACGTGATGTTATAACATCTCTCTCACACTGCCAGAACAGTCTTATACCTTGTCGGAGTGAAGGACATCCTTAACTAAGTAGATTCACTAAGCTATACTTAAAAAGCAATAAGGAATCATCTAAAATGTGTGACTCTTAACCCATATTGGCATACATGGTTTATGGGGGTTATTAATTGTCTGAACACTCCCCCATATAAATCAGTGATCAATATTAATCCCAATAATATACACTATTATGATTTGAGACTACACCCTGGAAGTGGCCGGCTCTCAAGAACCATTGCTGATCTCCAAGCCAAACCCTCATTCTGGTTGACTACAAGCTGAAGGCAAACTCAAGTATACAAAGCTTAAAACATAATAAAAATAATATACTCAACTCGCCACAAAATAGGCATTTAAGTCTTTAAAACATTTTTAAAAATAAATGAAACAAACTTCTCAAACTGTAATGTATATCTATGAAGCCTCTAAATGAAAAAAATGAAGGCAGATGAGACATACGGCATCCTAACAACTGATATAACTAAGAGTACAAGTGGAGCCCTTCGGATGTAAGGAGGCTCATCAAAGCTAATGTGAACTCCATGTGGTATCAATGAAGCACCTATTGATGACCGTGAATACATGTATCTGCATCATGAAACGATGCAGGCCAAAGGGCTTAGTACGTGAAATGTACGAGCATGTAAAGGGAATTCAAATACATAAACATAGGCTTGAACTTTGATATAAAGGAAACATACTTACCTATTTTTAACTCAAGAATAAAAAACATAGTTCAACTCAATGAAAAGACACTCAAGTCAGTGAAATAGGCCGCAACTCAATAATAAGATATTCGACTATGGGTAATCAACTCTGGGTACTCTATTCAATATAAAGTAAGAATACAAATGCATTATATGGAAAGACTTTAAAACGGTAGAAAACAACTCAATGTATTGAAAATTCAATAGTAAATTAGTTTGTATGTAAGGAACAATATAAACTTTGTTTGTATATGAAAATACAAAATAAACTTTGTGTATATAAAAGTACAAAATATCTCTGTGAAAGTTTCTCTAACCAACAACCATCACTATGAGCTTTCTGATAATACCACGTTTCGCCCATGATGTCAGAACTGTCCTATGATTTTCCAGTTCATAAGACCTACTCACTAAGTGGATCCACAAGTCTATGCTAAAAAATATTTAAGGAATCGTCTAAAAAGTATGACTCATTCTACCCACGTTGGCTACATGATTTATGGGGGTCGTAAGTTATCTAAACTCTCCTCCATATCGATGCGTAATGCTACTCACAAATATACTAGCTCACATGTTTAAAAATATAACTCGTTTTGTTTGAGATCATTACTCAAAATCCTTCTCTTAAAAGAGATGATACTCAAACTGCTCAAAACTCTTTTGGAAATCTCAAATTCGTCTCATCTTAAATGTAAAAATATTTACTCTTGGGAATACATAGTTATCATATATCATTTTAAAGAAAATGAACTCAACTCTGTTCTTTCTCAACTCAAGTGCTCAGTCTTAAACCAAATTAAAAAAAAGACTTCTCAAAATAAAGTTTATGTCGAATTATGGACGTGAACAATTCAATTCAAAGTTTTCGATAACCATAACTAAAACTAAATACTCGAGACTCAACATCTTAGAACTCAAGAACTTAAATGGTAATACTTCTTTCAAGAATGCTCGACTCAGAAGGTTAATGCAGAATAATGTGCATGAATTACTCAACTAAAGGACTCACTGATACTACTCAATCTCAAGATTGCTCGACTCGTAGGGTTAATGCAGAATTATGTGCATGAACTACTCAACTCAAAGACCTTCATAGGTAACATGTAGTAGCCCCATGATTTGGAATATAATCCCAAAATGATTAGGAACTCAATACTCAGGACTTAGAACTTGAAGATAATACTACTTCTCTCAAAGATACCCAACTGACGGAGTTCATGCAGAATTTATGGGCATGAACTACTCGACTCAAGAGTCTAAAACACAATATGACACTCATGTATATAACTCTTCTCATTCTAATACTTGTTTTCTCAAAACTCGGTTTAACTAAATAGTTGATCTCAAAGGATTCACAATTGAACTCAAAGACTTTCTTTGACTCCACTCTTAATTCTCTCTTAAATTTGTATTTGAATTATGAATTTAAGAGTTATGATTCATGATATGGGGAATCTCAATAACAATATAGAAATTTGATAATTAGGAATAGTACTTTTAAAAGAAAACATGAATTCAACTTAAAATCAACTTATCTAAAAAATATTCAAATATAGGGAAAGTATCCTAGACTACTGTGCTACTGATCTGAAAGTAGATGTAGGATGTGAGGATGAACTAGTCCAACACTATGATAGCCTTACATACCTGGAATAACGAGGTTCTTGGAAAATCTTCACTTGAAGAAGAACTTGATTAGAAGCCTTGAAACCTAGCTTGAAGGTAAACAATCAAGAAAACCTTTCTTAAGATTCTTGAATTAGTTTATGAAAATCTCTATGACCAAGCATTTTGATTTTCACTAGTGATTCATAATTGTATGGAGGAATTTGAATTGAAAAAGATGAAATGCTTGGAGAAAAGCTATCTTTGAAGAAGCTTGAAAAAGATTGGAAAGTCCTGTACTTTGATTTTCCCTTAGGATTTTGTCTTAGGGTTTGAGATAGAAAAGAATGATGGACTAAAAGATGAAAATCTAATTGTTTGGATCCTTTTTCAGCCAAGAAATCCGTTTAGGGTTTTCTTGGAGACAAACAAAATAAAAAAGACCATTTTTAATATTTTTCCGTCGGCTAATTCGTAATAACATTGTATCATGTTATTGAAAGAGTCATAACTTTTTACTCAAAAATTGGATTGATGCGAAATTAGTGGTGTTGGAAAGTAGATTCAAGTACCTCTAATTGGATAGGTTATTCCCTACATAAGTCTTTATATTCTAAAAGATATGGTTGTTTGCACTTGACCTAAGTAGAATTTTACATGAAAACTTAATAGAGAAGGAAACTTCAAGAACTCATCAAGAAATTTCAATTGCTCAATATTTATGGATAAATTTGTAGAAGAAACTCATGATTGACATGCGGGTGAATAAACCCAACACTATGGAAGCTTACATACCTCAAAGAACTAGGTTCTTGGCGAAATCTTGAATTTCTTCAACGAACGCTTGAAACTTTGAACTTTTTCTCTTCTTGAACTCTCAACTAAAACCCTAGGCGTATATTAGGATTATAAAAGTTAACATGATAGGATTAGACCTTTAAAAACTTTCTAAAATGAATTAAATCTGATTTAGCATGAAAAAGACCAAAATACCCCTTACTATTTTCGGATAACTTTTCTTAATTGGACTGCCTGACTTCAAAAAGGTATATCTCACTCATCCGACCTCAAAATTTAGCAAATTCAGTGGCGTTAGAAAGCTAATTTAAACACCTTTCATTTTCCATCTCATGGCACACATAACTCATTCTTTAAAGAGAGCTATGATCGTTCAAATTAACTCAAATCTTAGAAGAATTTAGGAATGTCTTGAACGAGCTACATCTAGTGACCTTAACACTTTGGAAAATTTTAAATTTCTTAGTAAAAACTTACTCACTATGAAGGATGGTTCAAGTCTTAGCTCAAAATTTTCCTAAGTTGCTATATATACTCATGCTCATATGTTTAAAACCAAAACCCTTCCTCGATTTGAATTAATTACCAAAAAGATTCTCTTAAAAAGATAATGCTCAAAACTCCCCCTAAACTCATTTGGAAATCTAGGTTTCCCTTGTTTTAAATATAAAAACATTTACTCTTGGAAATATTTAGTTCTCAGATATTCACTTGAAAAAAATTAAACTCGACTCTCATCATCTTCATACTCAAGTGCTCAAGTCCTAAAACAATTTATAACTAATTGTATAAGACTTCTCAAAATAGGGTTCATTCCGAATTATGGACGTGAACGACTCAATTCAAGGATTTCAATAACCATATATATAACTCAATAATAGGAACTCAACAACTCCAGAACTCAATGATACTACTCATCTCAAGAATGCTCGACTCACAGGGTCTTTGCGAAATTATTGGGCATGAACAACTCAACTCAAAGACCTTCATTTATACCATATGGTAGTCCCATAATAGGAATATAATCCCAAAAAAATTAGGAACTCAATACTCAAAAACTTAGAACTCGAAGATATTACTCATCTCAAAGATATTCAATTTATGGAATTCATGCTGAATTATGAGCATGAACGACTTGACTCAAGGATCTCAATAATAATGTAGACTCATGAATACACTCTTCTCATTCTCATACTCACATACTCGAGTATTAAAATAAATTATAAGTAATTGCAGAAGACTCCTTGAACAGACTCAAAAGGACTCCTTCGAATTTTACTCTTAATGCTACCTGAATTTTGTATTATAAATTTAAGGATCATGATTATGATATAAAGAATTTCTCAGCATATATGAAATGAACGAATTTGAGCATTGAACGTCTAACCTCATTTTTTAATTATTGTGATATGTAGAGTGGTGCAAAATCACAGATACCTCTCTTGATGCATTTCTATAGTTACGTTGATGTGAGATTATATATAGTTCAGCAGCAGCATGTTGGGAAAATTACTAATAACTCTTCTTTTATATCAAATTGTTGAAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGTTACCGGTTAGTGATACTCAGGTATTGTTTATCTACTTTATCATGTCGTAAGTATATTATTTGTAAAGATATATATCAAGATAGTTCGATTGCGTACACTTACATTTTGATTATGTTTGGTGAATACTATTCTAATACCTTTTTTTTTCCTAAAGCCTAACAAATAAAGATAATTAAGATGGGAACGTAATTCAAGTACAACATGGTTCCATACGTGACATATTTACACATATAGTGGAACCAAAAGAGCAATTTTTCCTAATATCATTTTCTAAATATCACGTGTGCCCGTGATTCTTTTTTATGGACATGAATTTTTTTTTTAATATGAGTGGAAGTAAGGTTCGATCTTTCTATCTGCTTTGATATCATATTGAATCGTGTGATTGTCTCTTTAAAAAATTAAGCAAGAGCATATTTTATTAATTAATTGTCTTTCTCGACGTTTTTCTCTTTCAACAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTCTTCAAAGGTAAGATATTAGTGATGTAATTAAATGATTTTAGTTAGATTTACATAAGTTTTTAATAAGTGAAAATTAATAGACATATTCTTGGAGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGGTATATCCAAATACTATAACTTAAATATATTGTAACGATTTAATTAATAGCATGTGTCACGTTCATCTATTCTTTAGTCACAATATATAGGGGCATGTCCTTAACAACGTGCCATGCCTCGATAGTCATTTTTGTCTTTTTGTGCGTATGAATTTAACTTTGACACAAATTTTTGTAGTAATAATAACTCATGCTTTAGCATCTTAGGAAGCAGTCATATGAAAAACAGAAGCATATATATATATTACATGAGTTAATTTAATTTAATATAAAATTTAATAAAATTGTGTCTCGCTATAAATAATTTTATTAAAAAATTATATAAATATATTATTTTTTTAACTGGCCGCAAAGTTATATAAATTGATAGAGAAAGAGGTTTTGGTGTAAGGTTCATTTTCCAACAATTAGTTTTATAATTTGTAAGTGCACACTTTATCAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGTACACTGCCTTAACATTACAAAATTAATTTATTTCATCAAAAGCATATCATAAAATTCTGACAAATAAATATATTAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGGTACATATCTATATATGTGTGTTAATTAATTAAGTTGATTTTGTATTTTTGTTTAATGAATAATTGTTTGTGATCATCAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGGTAAGTATCTTATTTTATATGACTTAGTTTGACTTGACATAAAGTTTAATAAAGAAAGAAAGACTTTTAAAACTTATAGTGTAAAATAAGTGAATAGATATATATGTGGTTGTACTAACACTACAACAAAAATAATTTTCAGCGGCATTAAATATTGACATTAATAATGAGTGCTAAAGACTTTATCGGTATTAGTTAAGTGTCATTAGGATCAATGTCGTTAAAGGCTTCACGGACATATACAAAGAGTGACAATTGCCGCTAATGATTATTTTTGTTGTAGTGAAAATGAGTATTTTAAAGTTAAATTGTTACATAATATAGAAATATGTCAGAAACAGGACAAATATACCACCGAACTATCATATATGTTATGGAGATATTCTCAGTCATACTTCTGCGACATTGGTACTCATGTCGTCCAAAAACTAGAACATATATATACCCTTTATATATTAACGAAGATACAAGTGTCATAATCTTATGCACCGATTCGATATTTATTAAATATCGAATCGACGGATAAAATTATGTCACGTGTCCCTATTAAGTCTTCTATTAGAGTAAAAAGCATATATTCTCTAGTTTTTGAACGAAAAAAGGTATTAATGTCTCAAAAGTATAACGAAAAGCATTTGCATACAATTTATGATAATTTGGGGCATATTAATTTATCATTCCCCCTTTTTTTGGCACTGATTAAAAAGAAAAAGAAAGTTATAAAAATTGGGATAGAGGGAATAATTGTTTCATAGGGAAAACTTAGAAGCTTCTCAGTATGTCAGTGAGAATGTGTTTCCTAATTAGTGAACTATGGTTTGGTGAAAAATAAAGAGAAAAAAATCAGTACAAATTTTCCACTGATTAGCAATGAGAAAAATATTTGTTTCTAGTAGTATGAGGAGAGGATAGTCCGCATAAATAATCCTTAAATTTGTGGATAAATAAACTATTTTCAATAGATTATCGTCTCAAAATAAAATAAAATGATTGCAAGAAAAGAATAATAGGTATGCTGGTAATATGTATAATACACTCAAATTTATTTGCTGTCCATGCAGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAA >allele-2 (SEQ ID NO: 11)ATGGGAAGAGGAAGAGTAGAACTAAAGAGAATAGAGAACAAAATAAACAGGCAAGTTACTTTTGCTAAGAGAAGAAATGGACTTCTTAAGAAAGCTTATGAGTTATCTATACTTTGTGATGCTGAAGTTGCTCTCATCATCTTCTCTAGCCGCGGAAAACTCTATGAGTTTTCAAGTGCTTCCAGGTATATATATATATACATATGTTTTTCTTCTTTTTGTGTGTGCGTATGTGTTTACTTACTTTCATTAATTAACTCAACCATATATATACATCTCTCACCTCAATTATATATATGTTTGAGATCTGAATGTCTACGGACTCCATTTAGGTACATATCTTTGTTTAGATCATAAATCATCTATCTTCATTCCTAAGATCTACTAATATATATGTATAAGAAGATCCATCCATCTATTAGGTTTTTCAACAACATATACAGTGAAATCTTATATGTGGGCCCACGTATAGCCATATGAGAAAATAGTGTGCACGTAAACATTATCATTACTTAATTATAGGAATATACATCCATTAGGTTTATCAACAACAATAAAATCCTCTAAATGGAGTCTAGTCATAGGTCTAGCCGTTTGAAAATGTAAAATATATGCCGATCTTATCACTATGTCATAATAATAGATATGTTGTTATTGAAAGATTCTCAATCTTTTTTTTTCTTCAAGGTAGAGATTCTTAAGTGGATTCATGTTTTTTTTATCAAAAAAGAAAAAAACAAAAGTGTCCATTTGTTCATCTAATGGGTTTTCCATGTTACCAATTCACTACACTGTTGAGATTTGATTATCAGATGTGTCAAGTTTCGTTTGGTTCCCTAGAAGGGAGAAAAGGCTGCTTATGCAGGCAGGGTATTAAAGATGATATTAATATCTGCAGTAATCAGTAACAGAATATATAAACTTAATAATAAACTTGAAGGTACTTAATTATCCAGCAGATAATCTTCTGTCTCACCGTACACTTTTGTTATATCATAAGCATAAGAATTGTTTTATCAAATATTACCAAACAAAACTTAGTTTTGTTTGGTAATATTTTATAAAATATGTTACCGAAAGTTACTTCCTATAACATATTTTATAAAGAAAAAAATTAAAAACTCCATATACCTAAGAAATGTAACCCCCCCTCCATAACAACAATTTAACAAAAATAAAAACCTACTTTTTTTGAATTTGGTAAATTAGTTTTCTATCCTTTTTAGTAACTTCCTTTCTTATTTTCTTTTTATATTGGTAAAGTTTAATATTACACATTATTTTAACATGTTATAATTTTTTGTGATGCTTAATTATTTGATACATGTAATAAACCATATATTAGAGCTATAAATCAATGACAATGCATGTAGATACAACTCATTTATGATATATTTTGTTTATATATATAACCAATTAGATAATTTGTCTGCGCTTTGTGCAGTCATAAATAATAATTGCATTGAACTTGCAAATATTTTTTTTTAATATCCATACATTAAAAAAAAAGAAAGAGGAAAATTGGTTCCTAAAATATTAGCAATATTCAAACATTTATTTGATTATTAATCATTATCACATAACTTAAGAACGTCTAATGAATGAATTATTCACGAAATAATAAATCATTGGTTCTAAAAAGGAATTTCGTAATAAAATAAAAATTTAAGTTACCATATTCAAAAAAAGAAATTGTGCTTGAACATGAAAATAATTATAATTTTTGAACTTGTATAATGAATTTCTTCAATTCATAAGTGGGAAATTTCATATTTATGTAATAATAGATAATATGTAAGCTCTAATATAGTACTTTAGGTTATAGAATTTAATATAAAATATCAAAACATGAATTCTTGAAATTGAGTAGAGTAATTATTTTCTGCACAATGAATCGGAGACAATAACTTTGAAGAAATATAAACAATAGAGTTCAAAAGATGTAGTCAAAAACAACAATTAATATCATAAGAATAAATTAATGAGTGTAAAAATGCATACCACGATATGTAAAAACAGAATGGAATATAATAAAAAAAATCGAGTTCACTGAATACACAATGTTCCTTTAAGAAAATTATTCTCCTCCAATACCAACGAGATTACATCCTCTAAGGATGGAAATGATTTCATTCCCCAACTTATCCATATAAAAATAGTGGTGTTAGTATGTAACTCAATAGGAGTAAAATACACAAATATTTAATTTTGCGAAAGTAGAAGAAGAAGATCATATTTTTTTTTTAAAATGAGAGGATATATCACTATTTTTAAACAACAAAGGGTAGTGTTAACAAATTTTTATTGTGTCTTGTCTAAAAGGTTACAGCTATTTGAAAAAGTTACAACACTTCGAAAAGTGAACAACATTTCATAAAAGTCGTAACTTTTCATAAAGTCGTAACTCTTCATAAATGTCGCAACTCTTCATAAAAATTACAACTATTGATAAAAGTCACCACTCTTGATAAAGATCACCACTCTTCATTGAAGTTGCAACTTTTCATAAAAATCACATCTTTTAATAAAAAAGAAAGACTAGTTTTTGGAATAAATTAATTTAAAAGAAAATTTTTGTTTGTGGTGGGGCGCCAAGTAGGCAGGCGTAGGGTTCTTTTTATATAAATATATATGATATATGATTCAATATTTGATATATATATATATAGAGAGAGAGATGACAATATAAGACAATTGCAAAAAATAAAATAAAAAACTAATCGAGTAAGTAGGCAAAAAATTATTTATAAAATATATGTAGAATTTCTTTATCAGATATGACTGCCCAAATCTTATATTCAAACTAAAATGCAAGATCAATGGTGCTATATATAGGGTTTTACACAAAAATCAAGATCTAGTCTTGCAAATTTAAATAAAAAACAGTGGTTTACGATGAGATAATGTAGCTTTTGTAAACAATAAAACTAGAAAAATAAATGCAAAGGCATTTTAAAGGATATAATAATGAAGATCAAAGGCAGAGAAGGGAAGAGGCAGCAATATAATGAAGGTAACATCATGGTTCCATTCTAATATATATGCTATTTTTCTTTAGTAAATTTCAAAAATAATGATACATTTTCATATTTGATAAATATTTAATGATACTATCAACATTTTATCTATATTGAGTTCCATTTATTTGACCAAAACCTCACAAAGATGTGCTCTTCGATCTATTCAAAATTTATTCAATTTAAGGATAGCTTTAAAACATGACAAAGTTTTCTCATATATTTCTTAAATTTTATATCCAGTCTAAATACGTATATAAACTAAAATGAAGAGAATAATATGAAGCTTTATTTGATGACATTGTTGAAATAACCAAAAGCTATAAGTGATACAATAGTAAATTTACCATTGGTCAATTCAGAATTATTTAAAAGCTAAAAAAGTCATATAAGTTGGGGTTGCTCAATGTATAGTTTTTGGCTTGTTTTAAGCATTTTAAAACTTTTTTTAAGCGCTTTTTAACATTGCTAAACACTCAAAAAATGATAAATAGTATTTAAATTTGATATGATTAGCTTAAAAGTGAACTCATATACCTTCAAAGTAAAAATCCCCAATTCGAGCTTTCAAACCACTTGATTTTGTGGATGAAATTATACTGAAGTTGAATATATCACTATTTATAGGGGTTAGTGAACTAATACCTTTGATTATTTGGTAGAAATATGTATCTTAGATCACCCTAATGAGCTCCCACTTTTAAAATAGGAAAAACCTCATATGAAGTTCATCACTGTTCATTATATATCACTTTTATTCAAAAACGTTTACAAATGTTCATTGTGACTAAATACCCTTGAGTGTCGAGTTTTCACACCAATAAGGCCTAATTAATAGGTAAACAAAACTATGTCAATCTTCAAAACGCAAATCTAATTATATTTTTAACAAGATTAGAGGTATATATACATATTCTCTTATGTTAACTCTTATTCATTATTGAACAAACTAAGTAAGTGTACCCAAGGTCTCAAACAACAGTTGGTACATTCTTTGTATGTCTTCCTTTGTCTCTTAATAGTCGTCTCCTCCTGTCGATGATTCCTCCAAATACATTAATCAAAGGAAAATCTTTCGCCCTCAACTTGCAAACTTGTCTATCTAAAATTGTTAACAAAGTTTCTTCATTAGAGAAACTATGATTTCTTGAATGTAGCAATTTGATGTGCCATGACTATCATCTTGATCAACATGCTTCTTAACCATCAAAAGATCCTAAACTAGATGCATGTCATGTTAGGAGACATATTAAGCTTGTATATAACTACACCAACATGCTTTAGGATCTCATAAGATCCAAAATTTCTTATTTGGGAGATTTTCAATCCAACAACCATCATAATGAGCAACGTGATGTTATAACATCTCTCTCACACTGCCAGAACAGTCTTATACCTTGTCGGAGTGAAGGACATCCTTAACTAAGTAGATTCACTAAGCTATACTTAAAAAGCAATAAGGAATCATCTAAAATGTGTGACTCTTAACCCATATTGGCATACATGGTTTATGGGGGTTATTAATTGTCTGAACACTCCCCCATATAAATCAGTGATCAATATTAATCCCAATAATATACACTATTATGATTTGAGACTACACCCTGGAAGTGGCCGGCTCTCAAGAACCATTGCTGATCTCCAAGCCAAACCCTCATTCTGGTTGACTACAAGCTGAAGGCAAACTCAAGTATACAAAGCTTAAAACATAATAAAAATAATATACTCAACTCGCCACAAAATAGGCATTTAAGTCTTTAAAACATTTTTAAAAATAAATGAAACAAACTTCTCAAACTGTAATGTATATCTATGAAGCCTCTAAATGAAAAAAATGAAGGCAGATGAGACATACGGCATCCTAACAACTGATATAACTAAGAGTACAAGTGGAGCCCTTCGGATGTAAGGAGGCTCATCAAAGCTAATGTGAACTCCATGTGGTATCAATGAAGCACCTATTGATGACCGTGAATACATGTATCTGCATCATGAAACGATGCAGGCCAAAGGGCTTAGTACGTGAAATGTACGAGCATGTAAAGGGAATTCAAATACATAAACATAGGCTTGAACTTTGATATAAAGGAAACATACTTACCTATTTTTAACTCAAGAATAAAAAACATAGTTCAACTCAATGAAAAGACACTCAAGTCAGTGAAATAGGCCGCAACTCAATAATAAGATATTCGACTATGGGTAATCAACTCTGGGTACTCTATTCAATATAAAGTAAGAATACAAATGCATTATATGGAAAGACTTTAAAACGGTAGAAAACAACTCAATGTATTGAAAATTCAATAGTAAATTAGTTTGTATGTAAGGAACAATATAAACTTTGTTTGTATATGAAAATACAAAATAAACTTTGTGTATATAAAAGTACAAAATATCTCTGTGAAAGTTTCTCTAACCAACAACCATCACTATGAGCTTTCTGATAATACCACGTTTCGCCCATGATGTCAGAACTGTCCTATGATTTTCCAGTTCATAAGACCTACTCACTAAGTGGATCCACAAGTCTATGCTAAAAAATATTTAAGGAATCGTCTAAAAAGTATGACTCATTCTACCCACGTTGGCTACATGATTTATGGGGGTCGTAAGTTATCTAAACTCTCCTCCATATCGATGCGTAATGCTACTCACAAATATACTAGCTCACATGTTTAAAAATATAACTCGTTTTGTTTGAGATCATTACTCAAAATCCTTCTCTTAAAAGAGATGATACTCAAACTGCTCAAAACTCTTTTGGAAATCTCAAATTCGTCTCATCTTAAATGTAAAAATATTTACTCTTGGGAATACATAGTTATCATATATCATTTTAAAGAAAATGAACTCAACTCTGTTCTTTCTCAACTCAAGTGCTCAGTCTTAAACCAAATTAAAAAAAAGACTTCTCAAAATAAAGTTTATGTCGAATTATGGACGTGAACAATTCAATTCAAAGTTTTCGATAACCATAACTAAAACTAAATACTCGAGACTCAACATCTTAGAACTCAAGAACTTAAATGGTAATACTTCTTTCAAGAATGCTCGACTCAGAAGGTTAATGCAGAATAATGTGCATGAATTACTCAACTAAAGGACTCACTGATACTACTCAATCTCAAGATTGCTCGACTCGTAGGGTTAATGCAGAATTATGTGCATGAACTACTCAACTCAAAGACCTTCATAGGTAACATGTAGTAGCCCCATGATTTGGAATATAATCCCAAAATGATTAGGAACTCAATACTCAGGACTTAGAACTTGAAGATAATACTACTTCTCTCAAAGATACCCAACTGACGGAGTTCATGCAGAATTTATGGGCATGAACTACTCGACTCAAGAGTCTAAAACACAATATGACACTCATGTATATAACTCTTCTCATTCTAATACTTGTTTTCTCAAAACTCGGTTTAACTAAATAGTTGATCTCAAAGGATTCACAATTGAACTCAAAGACTTTCTTTGACTCCACTCTTAATTCTCTCTTAAATTTGTATTTGAATTATGAATTTAAGAGTTATGATTCATGATATGGGGAATCTCAATAACAATATAGAAATTTGATAATTAGGAATAGTACTTTTAAAAGAAAACATGAATTCAACTTAAAATCAACTTATCTAAAAAATATTCAAATATAGGGAAAGTATCCTAGACTACTGTGCTACTGATCTGAAAGTAGATGTAGGATGTGAGGATGAACTAGTCCAACACTATGATAGCCTTACATACCTGGAATAACGAGGTTCTTGGAAAATCTTCACTTGAAGAAGAACTTGATTAGAAGCCTTGAAACCTAGCTTGAAGGTAAACAATCAAGAAAACCTTTCTTAAGATTCTTGAATTAGTTTATGAAAATCTCTATGACCAAGCATTTTGATTTTCACTAGTGATTCATAATTGTATGGAGGAATTTGAATTGAAAAAGATGAAATGCTTGGAGAAAAGCTATCTTTGAAGAAGCTTGAAAAAGATTGGAAAGTCCTGTACTTTGATTTTCCCTTAGGATTTTGTCTTAGGGTTTGAGATAGAAAAGAATGATGGACTAAAAGATGAAAATCTAATTGTTTGGATCCTTTTTCAGCCAAGAAATCCGTTTAGGGTTTTCTTGGAGACAAACAAAATAAAAAAGACCATTTTTAATATTTTTCCGTCGGCTAATTCGTAATAACATTGTATCATGTTATTGAAAGAGTCATAACTTTTTACTCAAAAATTGGATTGATGCGAAATTAGTGGTGTTGGAAAGTAGATTCAAGTACCTCTAATTGGATAGGTTATTCCCTACATAAGTCTTTATATTCTAAAAGATATGGTTGTTTGCACTTGACCTAAGTAGAATTTTACATGAAAACTTAATAGAGAAGGAAACTTCAAGAACTCATCAAGAAATTTCAATTGCTCAATATTTATGGATAAATTTGTAGAAGAAACTCATGATTGACATGCGGGTGAATAAACCCAACACTATGGAAGCTTACATACCTCAAAGAACTAGGTTCTTGGCGAAATCTTGAATTTCTTCAACGAACGCTTGAAACTTTGAACTTTTTCTCTTCTTGAACTCTCAACTAAAACCCTAGGCGTATATTAGGATTATAAAAGTTAACATGATAGGATTAGACCTTTAAAAACTTTCTAAAATGAATTAAATCTGATTTAGCATGAAAAAGACCAAAATACCCCTTACTATTTTCGGATAACTTTTCTTAATTGGACTGCCTGACTTCAAAAAGGTATATCTCACTCATCCGACCTCAAAATTTAGCAAATTCAGTGGCGTTAGAAAGCTAATTTAAACACCTTTCATTTTCCATCTCATGGCACACATAACTCATTCTTTAAAGAGAGCTATGATCGTTCAAATTAACTCAAATCTTAGAAGAATTTAGGAATGTCTTGAACGAGCTACATCTAGTGACCTTAACACTTTGGAAAATTTTAAATTTCTTAGTAAAAACTTACTCACTATGAAGGATGGTTCAAGTCTTAGCTCAAAATTTTCCTAAGTTGCTATATATACTCATGCTCATATGTTTAAAACCAAAACCCTTCCTCGATTTGAATTAATTACCAAAAAGATTCTCTTAAAAAGATAATGCTCAAAACTCCCCCTAAACTCATTTGGAAATCTAGGTTTCCCTTGTTTTAAATATAAAAACATTTACTCTTGGAAATATTTAGTTCTCAGATATTCACTTGAAAAAAATTAAACTCGACTCTCATCATCTTCATACTCAAGTGCTCAAGTCCTAAAACAATTTATAACTAATTGTATAAGACTTCTCAAAATAGGGTTCATTCCGAATTATGGACGTGAACGACTCAATTCAAGGATTTCAATAACCATATATATAACTCAATAATAGGAACTCAACAACTCCAGAACTCAATGATACTACTCATCTCAAGAATGCTCGACTCACAGGGTCTTTGCGAAATTATTGGGCATGAACAACTCAACTCAAAGACCTTCATTTATACCATATGGTAGTCCCATAATAGGAATATAATCCCAAAAAAATTAGGAACTCAATACTCAAAAACTTAGAACTCGAAGATATTACTCATCTCAAAGATATTCAATTTATGGAATTCATGCTGAATTATGAGCATGAACGACTTGACTCAAGGATCTCAATAATAATGTAGACTCATGAATACACTCTTCTCATTCTCATACTCACATACTCGAGTATTAAAATAAATTATAAGTAATTGCAGAAGACTCCTTGAACAGACTCAAAAGGACTCCTTCGAATTTTACTCTTAATGCTACCTGAATTTTGTATTATAAATTTAAGGATCATGATTATGATATAAAGAATTTCTCAGCATATATGAAATGAACGAATTTGAGCATTGAACGTCTAACCTCATTTTTTAATTATTGTGATATGTAGAGTGGTGCAAAATCACAGATACCTCTCTTGATGCATTTCTATAGTTACGTTGATGTGAGATTATATATAGTTCAGCAGCAGCATGTTGGGAAAATTACTAATAACTCTTCTTTTATATCAAATTGTTGAAGCATGATGACAACACTTGAAAAGTATCAACAATGCAGTTACGCATCTTTGGACCCGATGTTACCGGTTAGTGATACTCAGGTATTGTTTATCTACTTTATCATGTCGTAAGTATATTATTTGTAAAGATATATATCAAGATAGTTCGATTGCGTACACTTACATTTTGATTATGTTTGGTGAATACTATTCTAATACCTTTTTTTTTCCTAAAGCCTAACAAATAAAGATAATTAAGATGGGAACGTAATTCAAGTACAACATGGTTCCATACGTGACATATTTACACATATAGTGGAACCAAAAGAGCAATTTTTCCTAATATCATTTTCTAAATATCACGTGTGCCCGTGATTCTTTTTTATGGACATGAATTTTTTTTTTAATATGAGTGGAAGTAAGGTTCGATCTTTCTATCTGCTTTGATATCATATTGAATCGTGTGATTGTCTCTTTAAAAAATTAAGCAAGAGCATATTTTATTAATTAATTGTCTTTCTCGACGTTTTTCTCTTTCAACAGATGAACTACAATGAGTATGTGAGGCTAAAAGCTAGAGTTGAGCTCCTTCAACGTTC

TCAAAGGTAAGATATTAGTGATGTAATTAAATGATTTTAGTTAGATTTACATAAGTTTTTAATAAGTGAAAATTAATAGACATATTCTTGGAGAGATTTGGGCACACTAAACTCGAAAGAACTTGAGCAGCTTGAGCACCAATTGGATGCATCTTTGAAGAAAGTTAGATCAAAAAAGGTATATCCAAATACTATAACTTAAATATATTGTAACGATTTAATTAATAGCATGTGTCACGTTCATCTATTCTTTAGTCACAATATATAGGGGCATGTCCTTAACAACGTGCCATGCCTCGATAGTCATTTTTGTCTTTTTGTGCGTATGAATTTAACTTTGACACAAATTTTTGTAGTAATAATAACTCATGCTTTAGCATCTTAGGAAGCAGTCATATGAAAAACAGAAGCATATATATATATTACATGAGTTAATTTAATTTAATATAAAATTTAATAAAATTGTGTCTCGCTATAAATAATTTTATTAAAAAATTATATAAATATATTATTTTTTTAACTGGCCGCAAAGTTATATAAATTGATAGAGAAAGAGGTTTTGGTGTAAGGTTCATTTTCCAACAATTAGTTTTATAATTTGTAAGTGCACACTTTATCAGACTCAATCTATGCTGGATCAGCTGGCAGACCTTCAAGAAAAGGTACACTGCCTTAACATTACAAAATTAATTTATTTCATCAAAAGCATATCATAAAATTCTGACAAATAAATATATTAGGAGCAAATGCTGGAAGAAGCAAATAAACAACTAAAAAACAAGGTACATATCTATATATGTGTGTTAATTAATTAAGTTGATTTTGTATTTTTGTTTAATGAATAATTGTTTGTGATCATCAGCTGGAAGAAAGTGCAGCTAGAATTCCACTTGGATTGTCATGGGGAAATAATGGAGGACAAACAATGGAATACAATCGACTCCCTCCACAAACTACTGCACAACCTTTCTTTCAACCTCTCCGTTTGAATTCTTCATCGCCTCAATTCGGGTAAGTATCTTATTTTATATGACTTAGTTTGACTTGACATAAAGTTTAATAAAGAAAGAAAGACTTTTAAAACTTATAGTGTAAAATAAGTGAATAGATATATATGTGGTTGTACTAACACTACAACAAAAATAATTTTCAGCGGCATTAAATATTGACATTAATAATGAGTGCTAAAGACTTTATCGGTATTAGTTAAGTGTCATTAGGATCAATGTCGTTAAAGGCTTCACGGACATATACAAAGAGTGACAATTGCCGCTAATGATTATTTTTGTTGTAGTGAAAATGAGTATTTTAAAGTTAAATTGTTACATAATATAGAAATATGTCAGAAACAGGACAAATATACCACCGAACTATCATATATGTTATGGAGATATTCTCAGTCATACTTCTGCGACATTGGTACTCATGTCGTCCAAAAACTAGAACATATATATACCCTTTATATATTAACGAAGATACAAGTGTCATAATCTTATGCACCGATTCGATATTTATTAAATATCGAATCGACGGATAAAATTATGTCACGTGTCCCTATTAAGTCTTCTATTAGAGTAAAAAGCATATATTCTCTAGTTTTTGAACGAAAAAAGGTATTAATGTCTCAAAAGTATAACGAAAAGCATTTGCATACAATTTATGATAATTTGGGGCATATTAATTTATCATTCCCCCTTTTTTTGGCACTGATTAAAAAGAAAAAGAAAGTTATAAAAATTGGGATAGAGGGAATAATTGTTTCATAGGGAAAACTTAGAAGCTTCTCAGTATGTCAGTGAGAATGTGTTTCCTAATTAGTGAACTATGGTTTGGTGAAAAATAAAGAGAAAAAAATCAGTACAAATTTTCCACTGATTAGCAATGAGAAAAATATTTGTTTCTAGTAGTATGAGGAGAGGATAGTCCGCATAAATAATCCTTAAATTTGTGGATAAATAAACTATTTTCAATAGATTATCGTCTCAAAATAAAATAAAATGATTGCAAGAAAAGAATAATAGGTATGCTGGTAATATGTATAATACACTCAAATTTATTTGCTGTCCATGCAGATACAATCCAAATATGGGTGCAAATGATCATGAGGTTAATGCAGCAACAACTGCTCATAATATTAATGGATTTATTCCAGGGTGGATGCTCTAAWild-type Solyc03g114840 gene (SEQ ID NO: 12)ATGGGAAGAGGAAGAGTTGAGCTTAAGAGAATAGAAAATAAAATAAATAGGCAAGTCACTTTTGCTAAGAGAAGAAATGGACTTCTTAAAAAAGCTTATGAACTTTCTGTTCTTTGTGATGCTGAAGTTGCCCTTATAATCTTCTCTAATAGGGGTAAACTCTATGAATTTTGCAGCACTTCAAGGTATTTTTTATTTTATTATATTAACATCAAAGATTTTATTTTTTTAAAAAAAACCTTAAGTCCTTCATTACCAAAACCCTTAATTGATTTACAAAGTACTTTCATTAAATTTAGTAATTCTTTTTTTTTTTATCTCTGACTTCAATTATAATGCAAGATCTATGTTGTCTTTATATATATTGAATTATATATGTACTGTATTTTTACTATATACATATAAGATCCTTTTTTCTTTTTTTTCTGTCTCTTTATATAAATATATTTTAAATAGTTGATTTTGAAAGATCTACTAATGTATATTTATTTTTGGAACTTTTGTGTATATGGAATTTTTTTCTTTTTTATGTTTTTTTTTTGTTCTAATTGTTTTAAAAGCGTTTAAGATCAGAATGTTCTTGATTATTCTTTTAGGAAAAAGATTTCCCATACATTGAGTTATTTTTTGATCTGTAGATTGAATTTTTTTAATGAGTTCCGATAGATTTTCGTTCAATTTTTCAATGAAACTATTGAGGGTTGATGATTAGATAATTACTCGATTGAAAGTTTTTATTTCAAAAAAATTATAATTCTTCTTAATTTTATATTTATGAGATAGAGTTAGTTTAGTGATTATATGAAAAATCGTATCAGATTATTGGGAATCGAAACTTAAAAATTCTGAAAATATTATTATAAATTTTACATGTTACAATATTTTTACTGTTAAGATTTGATTTGCAGACTAGGTGTCATGTTTGACAGTTGATAAAAAATCTGTTATTTTTGTTCTTTAATTCCCAAGACGGATAAACAAAGGCTGCTTATGTTGGTTTCCAATAAGCAGCCATAATTTTAAATATTTTTGTTAAGATTAATTAATAACAATTATTTCCACCAGATAATTTTCAAAATTTGTGACCCCGAGTTCATATAAATTGTTAATTTTACTGCTAGAAATTACATCGATAATAATTTATTTAGTGTAATCTTATAAATACGAGGGCAGTAGTGTATAGACTGTTTTTTATTAATCCTGACTCAAAGTGAGGTAAGTTAAGTATATTTTGATTAAAAGGACTACATTTCATTTATGTATGTTTAATTAATATTATTTTGTAAGTCAATAAATCTAAACAACATGAGTTTATCTAGACCCTTAATTATGCACCTTCATTATCAATTTTTTCAATACTCTCCTCAGAACATATGCTTCTCTATAATTTTGTGCACGAGTTAATCAATTCTTCCTTTTCAATAATTAAATATGTGATTTATGTTTAGCACTTATTTTTCGGTTAGTTAATTGATAATAGGAAAAAGCCTCTTTTTTTTTGTGTGTGTGGTAATTAGGATCTTTATTGAATTTAAAATGACCTACTATAGAACTTGGGAGTTTTTCTTCATAATAATGCACTGCAACGTGTTAAAAAAAAAGAATCAAATGAAATTAATAGATGTTTACTGGATTGCCATGGTAAAGTGATAAGTATTAATTTCGCTTTAACTAAGAGATCATTATATTCAAGTCCCCTTGATACAAACTTGCCTTTGTAAATAAGTGTTTTATTTTTCAATGTGAAACTTTCGCTGTTAATTTAAATTTAATTATACTTCTATATAAATACCAAACAATAATGTAATAAAACAAAAAATAAAAGAGTAGATGTTTCATATTGTTAATGCAGCATGGTGAAAACAATTGAAAAGTACCAACGTTGCAGCTATGCTACTTTGGAAGCCAACCAATCAGTTACTGATACTCAGGTACTGCTTTATATTTTAATTTATTTGGCTTTTTTTTAAAAAAATAATTAGTTTTGATTAATATGCATCATTTTATTTATTTTTGGCAACTCTTTATTTATCAGTAATAAGTAATAACTTTTTAACTAGTATATTTAAAAATCACAAAATTTAAGAATATTTTAATAGATTCGACATATTTTAGTTTAAAAATAACAAATTAAATTATGTTTTTAATTTTTTAAATATTCTTACTATAATTATCATGTACTCTTTGATCTGTTCATCTTTTCCATGATAATATTATTTGGTCAGTTAGTGACATAAGAGTTTGAAATTTAGAAAAAAGGAATATTTGGAGAAAACTGAAATGGATATTTAGAAATGAAAGTTATTTAATATAAATATAAGTATGGGCTGCTGAGTTGGGAATCCACGCTGGAGATCTCAAGTTTGAAGCGTCTCACAAACAATAGTAATGTCTTTTTGGTCGAGTTTGTCGGATTGGACTTGTCCGTGGCCTGTGGGTTACTTTTCCTATATGGTTTGCAAGCTATCGGGAATTTTATCCTGGCGCACCCAAATTTGAGTTATTTTTGAGTTTTTATATGAAATAGCTTTGTGAATTCATCGAACTCCCGAAAACATTGAACTTTACTCCAAGTTGAATTGCAGTAAAATAATAGTAGCGATTCTTTAATTTATCCTAACAGTTTTTCGAAATAATAATCCCAAAAAAGTTTAAAATAACCATACCATAAACTTACTGGGTAAGATATTATCTGTCTAATAATATATAGTAGTTTCTTTTGTTTTATTAGTTTATCTAATCCATATTTCATTTCTTGATAAGTTATTCTTAATAGGAAAATAAACTTATTTCGAAAAACTGTTTTTAAAATTTTCTTGAGTTGAGTCTTGGATGAAAAATAGTTAATTTTGCATTAATTAATTTTGTTCTAACAAAAACTAATTAAATTTTTTTGAAGCGCATATTCACTCAAAAAATAAATAAAAACCATCATGCATACAGGAAATGTTCTTTTTTTAATTTATTTTTTCATTGGAGCCCTGACTAATTTTATATCGGTTCATACTTTCATAAATTACAAAAAGTTCAAAATTTAAACTAACCATATAAGTGAATAAAATAAATCAACAAAATATTCACCACATAATACTTTTTAAATAGAATTTTTCATACCAAAGACCTTACTTTAATTAATTAGGGTGAGAGAATCCTATAAGTCAATGCAAAACAATTCTATCTATCGGATTATAATCGTTGATTCATAAAATTTTAAAATCGACGATTTTCATTTAAATGACCCTTTTTTTTCTTTCATTTTTTATTGTTATTCATCTATTTAACTTGTGAGCATCTTTCATATTGATATTTCAGACCCTTAAATTAATTGTTTTCTTACAGAATAACTACCACGAATATCTGAGGCTAAAAGCTAGAGTTGAGCTCCTCCAACGATCTCAGAGGTAATTTCTGTTCACTATCTTTATCTCAAATGAATTCTCATGTTTTTATTTTTCGAGATTCAGATTAAATATAATTTGATGTATTATTAATTTAAATACGTTATTTAATATGGTCCTTATGTCCAACCATTGATTTAATTTGATATTTTTTTAATGAAAATTACACAGAAACTTTCTTGGTGAAGATTTGGGCACGTTAAGCTCGAAGGACCTTGAGCAGCTTGAGAATCAATTAGAGTCTTCCTTAAAGCAAATCAGGTCAAGGAAGGTAAATTATTTAATCTAATTATACAGAAAAATCATCTAAAAGTTACCTTAATTGCTAGCCCAATAAGTTTGCTATCTGTTGATCCTCACATTATTTTACTCACAGAAATTCACAATACCTTTATTTTTGTTTGAGTTTGAAGTATACAATTTCTTTAAAATGTAAAATTTGAAATCTCAACAATAAGATATGTTATTGATCCTTGCAATTATGGGTAGATTGCGAATTAAACTATCTTGTCTTTGCTTACAACAGTCATTTTGTTTATAAACTAATTATACATAAATCCTAACTGATAGATAGTTTATAAAGATGAATAATGAACATAGGTCATATATTAAAAAAACAAAAAACAAAAAAAAACTAAACAAGATGAGCGAGTCAAAAATAGTCTTAACAAAAGAATATATATATATGTATATATCATATTTGATTTGTCTATTTTTAATTTTGAAAAAACTAAGTTAATCGATATATAATATGAAGGCATAATGCATAAATATGTCCTTTAACTTGGTTTTAAATCACATTTATACCTCTTCGACTTTGGGTGTATACAAACAAACACTTAAACTTATATAATGTTGAACAAATAGATATATATGTCCTACATGTCATTTTTCGTCCTAAATGGTGTCCTAAGTGTATTGTGTCACGCAGGACTCATGTGTCTATTTGTTCAAATTTATACAAGTTTAAGTGCTTACTTATGTATAAACAAAGTTGAATGACATAAATGTGAAATAAAATCAAATTAAAGGGCATATTTATGCATTATACCTAATACGAAAATCCATATTATTCACTAAAAAATGAGTCGGATTATATGATTACTTTTTTATTCATTTTGCCAATCGTATCCTACGACATTGTTTTTAATTTGCAGACACAATTCATGCTGGATCAGCTTGCAGATCTTCAACAAAAGGTAATTATAAAATTCTACAAATTTCCAATAATTAATAAATGGAATAATTATGCGCGAGAAATTTATCTATTTAAAATTTACGATGAATTTTAATTTTACAGGAGCAAATGCTTGCAGAATCTAATAGATTACTCCGTAGAAAGGTAAACTAACTTGATAGCCGTGCGTAATGAATAACTTATTTTATTTTCAAAATTATAAATCTAAATACTTAGGTAACTCGATAACATAAGAAGTATTTATACTGATGATATTGGTGTTGTGTTTTTTTTTATTAGTTAGAAGAAAGTGTAGCTGGATTTCCACTTCGATTGTGTTGGGAAGATGGAGGTGATCATCAACTTATGCATCAACAAAATCGTCTCCCTAACACAGAGGGTTTCTTTCAGCCTCTTGGATTGCATTCTTCTTCTCCACATTTTGGGTAATTACTTTTATTATTATTAAAAATAATTTCAATTTTTTTTACTTTTATTTCGATTAATAAATCAATGTGCACCAAGGTACGGTCTAACATAAACAAAAATGTGGGGAATGCTCTTAAAGCCCTAACAAAAGTTATTTGGTACGTGTACTAATGTAATCGTACTATATATCTTACTTGATTAGTGGATGGACAGTACTGGGCACACACAATTGACATAAGTTATTATAAGGAAAAAAAAAGGCCAATAATCAATATAGTCCAACATTACATTATTTATTATAACAGGTCACTCTAGATTAAATGTTAATGAATAACAAAAAGTCTCATATTGATGATTAATGTGATGGGTGGGCTTCTTATAAGGCTTTGACAATCCTACTCTCTTTGAGCTAGTTTTGGGGGTGTGACCTAATTCAACAGAACGTAGTTAAGATTGTGAAGTAAAGTTGATCATTGTTATAACAGGTTTAAATACTTCTAGTAAAAATAGTTCCTAGATAATCCATCGCAAAATAGCTCCTATATAGTTAGTTGGATTTTCATATAATCTATAGCTTATACATAGCTAAATGGGAATAGATGAGAGTTTCTGTTGTTTAGATATGATATTTGATCGGTTTCTAAATCGTTACTATCATGTAGTGAATAATTTTCATGTTATTACTATTACATTTGATTGTTTCTGTGGTTATTTTTTTTTCTAGGTACAATCCTGTTAATACAGATGAGGTGAATGCAGCGGCAACTGCACACAATATGAATGGATTTATTCATGGATGGATGCTTTAAWild-type Solyc03g114840 coding sequence (SEQ ID NO: 13)ATGGGAAGAGGAAGAGTTGAGCTTAAGAGAATAGAAAATAAAATAAATAGGCAAGTCACTTTTGCTAAGAGAAGAAATGGACTTCTTAAAAAAGCTTATGAACTTTCTGTTCTTTGTGATGCTGAAGTTGCCCTTATAATCTTCTCTAATAGGGGTAAACTCTATGAATTTTGCAGCACTTCAAGCATGGTGAAAACAATTGAAAAGTACCAACGTTGCAGCTATGCTACTTTGGAAGCCAACCAATCAGTTACTGATACTCAGAATAACTACCACGAATATCTGAGGCTAAAAGCTAGAGTTGAGCTCCTCCAACGATCTCAGAGAAACTTTCTTGGTGAAGATTTGGGCACGTTAAGCTCGAAGGACCTTGAGCAGCTTGAGAATCAATTAGAGTCTTCCTTAAAGCAAATCAGGTCAAGGAAGACACAATTCATGCTGGATCAGCTTGCAGATCTTCAACAAAAGGAGCAAATGCTTGCAGAATCTAATAGATTACTCCGTAGAAAGTTAGAAGAAAGTGTAGCTGGATTTCCACTTCGATTGTGTTGGGAAGATGGAGGTGATCATCAACTTATGCATCAACAAAATCGTCTCCCTAACACAGAGGGTTTCTTTCAGCCTCTTGGATTGCATTCTTCTTCTCCACATTTTGGGTACAATCCTGTTAATACAGATGAGGTGAATGCAGCGGCAACTGCACACAATATGAATGGATTTATTCATGGATGGATGCTTTAAMutant Solyc03g114840 gene allele ej2^(W) (SEQ ID NO: 14)ATGGGAAGAGGAAGAGTTGAGCTTAAGAGAATAGAAAATAAAATAAATAGGCAAGTCACTTTTGCTAAGAGAAGAAATGGACTTCTTAAAAAAGCTTATGAACTTTCTGTTCTTTGTGATGCTGAAGTTGCCCTTATAATCTTCTCTAATAGGGGTAAACTCTATGAATTTTGCAGCACTTCAAGGTATTTTTTATTTTATTATATTAACATCAAAGATTTTATTTTTTTAAAAAAAACCTTAAGTCCTTCATTACCAAAACCCTTAATTGATTTACAAAGTACTTTCATTAAATTTAGTAATTCTTTTTTTTTTTATCTCTGACTTCAATTATAATGCAAGATCTATGTTGTCTTTATATATATTGAATTATATATGTACTGTATTTTTACTATATACATATAAGATCCTTTTTTCTTTTTTTTCTGTCTCTTTATATAAATATATTTTAAATAGTTGATTTTGAAAGATCTACTAATGTATATTTATTTTTGGAACTTTTGTGTATATGGAATTTTTTTCTTTTTTATGTTTTTTTTTTGTTCTAATTGTTTTAAAAGCGTTTAAGATCAGAATGTTCTTGATTATTCTTTTAGGAAAAAGATTTCCCATACATTGAGTTATTTTTTGATCTGTAGATTGAATTTTTTTAATGAGTTCCGATAGATTTTCGTTCAATTTTTCAATGAAACTATTGAGGGTTGATGATTAGATAATTACTCGATTGAAAGTTTTTATTTCAAAAAAATTATAATTCTTCTTAATTTTATATTTATGAGATAGAGTTAGTTTAGTGATTATATGAAAAATCGTATCAGATTATTGGGAATCGAAACTTAAAAATTCTGAAAATATTATTATAAATTTTACATGTTACAATATTTTTACTGTTAAGATTTGATTTGCAGACTAGGTGTCATGTTTGACAGTTGATAAAAAATCTGTTATTTTTGTTCTTTAATTCCCAAGACGGATAAACAAAGGCTGCTTATGTTGGTTTCCAATAAGCAGCCATAATTTTAAATATTTTTGTTAAGATTAATTAATAACAATTATTTCCACCAGATAATTTTCAAAATTTGTGACCCCGAGTTCATATAAATTGTTAATTTTACTGCTAGAAATTACATCGATAATAATTTATTTAGTGTAATCTTATAAATACGAGGGCAGTAGTGTATAGACTGTTTTTTATTAATCCTGACTCAAAGTGAGGTAAGTTAAGTATATTTTGATTAAAAGGACTACATTTCATTTATGTATGTTTAATTAATATTATTTTGTAAGTCAATAAATCTAAACAACATGAGTTTATCTAGACCCTTAATTATGCACCTTCATTATCAATTTTTTCAATACTCTCCTCAGAACATATGCTTCTCTATAATTTTGTGCACGAGTTAATCAATTCTTCCTTTTCAATAATTAAATATGTGATTTATGTTTAGCACTTATTTTTCGGTTAGTTAATTGATAATAGGAAAAAGCCTCTTTTTTTTTGTGTGTGTGGTAATTAGGATCTTTATTGAATTTAAAATGACCTACTATAGAACTTGGGAGTTTTTCTTCATAATAATGCACTGCAACGTGTTAAAAAAAAAGAATCAAATGAAATTAATAGATGTTTACTGGATTGCCATGGTAAAGTGATAAGTATTAATTTCGCTTTAACTAAGAGATCATTATATTCAAGTCCCCTTGATACAAACTTGCCTTTGTAAATAAGTGTTTTATTTTTCAATGTGAAACTTTCGCTGTTAATTTAAATTTAATTATACTTCTATATAAATACCAAACAATAATGTAATAAAACAAAAAATAAAAGAGTAGATGTTTCATATTGTTAATGCAGCATGGTGAAAACAATTGAAAAGTACCAACGTTGCAGCTATGCTACTTTGGAAGCCAACCAATCAGTTACTGATACTCAGGTACTGCTTTATATTTTAATTTATTTGGCTTTTTTTTAAAAAAATAATTAGTTTTGATTAATATGCATCATTTTATTTATTTTTGGCAACTCTTTATTTATCAGTAATAAGTAATAACTTTTTAACTAGTATATTTAAAAATCACAAAATTTAAGAATATTTTAATAGATTCGACATATTTTAGTTTAAAAATAACAAATTAAATTATGTTTTTAATTTTTTAAATATTCTTACTATAATTATCATGTACTCTTTGATCTGTTCATCTTTTCCATGATAATATTATTTGGTCAGTTAGTGACATAAGAGTTTGAAATTTAGAAAAAAGGAATATTTGGAGAAAACTGAAATGGATATTTAGAAATGAAAGTTATTTAATATAAATATAAGTATGGGCTGCTGAGTTGGGAATCCACGCTGGAGATCTCAAGTTTGAAGCGTCTCACAAACAATAGTAATGTCTTTTTGGTCGAGTTTGTCGGATTGGACTTGTCCGTGGCCTGTGGGTTACTTTTCCTATATGGTTTGCAAGCTATCGGGAATTTTATCCTGGCGCACCCAAATTTGAGTTATTTTTGAGTTTTTATATGAAATAGCTTTGTGAATTCATCGAACTCCCGAAAACATTGAACTTTACTCCAAGTTGAATTGCAGTAAAATAATAGTAGCGATTCTTTAATTTATCCTAACAGTTTTTCGAAATAATAATCCCAAAAAAGTTTAAAATAACCATACCATAAACTTACTGGGTAAGATATTATCTGTCTAATAATATATAGTAGTTTCTTTTGTTTTATTAGTTTATCTAATCCATATTTCATTTCTTGATAAGTTATTCTTAATAGGAAAATAAACTTATTTCGAAAAACTGTTTTTAAAATTTTCTTGAGTTGAGTCTTGGATGAAAAATAGTTAATTTTGCATTAATTAATTTTGTTCTAACAAAAACTAATTAAATTTTTTTGAAGCGCATATTCACTCAAAAAATAAATAAAAACCATCATGCATACAGGAAATGTTCTTTTTTTAATTTATTTTTTCATTGGAGCCCTGACTAATTTTATATCGGTTCATACTTTCATAAATTACAAAAAGTTCAAAATTTAAACTAACCATATAAGTGAATAAAATAAATCAACAAAATATTCACCACATAATACTTTTTAAATAGAATTTTTCATACCAAAGACCTTACTTTAATTAATTAGGGTGAGAGAATCCTATAAGTCAATGCAAAACAATTCTATCTATCGGATTATAATCGTTGATTCATAAAATTTTAAAATCGACGATTTTCATTTAAATGACCCTTTTTTTTCTTTCATTTTTTATTGTTATTCATCTATTTAACTTGTGAGCATCTTTCATATTGATATTTCAGACCCTTAAATTAATTGTTTTCTTACAGAATAACTACCACGAATATCTGAGGCTAAAAGCTAGAGTTGAGCTCCTCCAACGATCTCAGAGGTAATTTCTGTTCACTATCTTTATCTCAAATGAATTCTCATGTTTTTATTTTTCGAGATTCAGATTAAATATAATTTGATGTATTATTAATTTAAATACGTTATTTAATATGGTCCTTATGTCCAACCATTGATTTAATTTGATATTTTTTTAATGAAAATTACACAGAAACTTTCTTGGTGAAGATTTGGGCACGTTAAGCTCGAAGGACCTTGAGCAGCTTGAGAATCAATTAGAGTCTTCCTTAAAGCAAATCAGGTCAAGGAAGGTAAATTATTTAATCTAATTATACAGAAAAATCATCTAAAAGTTACCTTAATTGCTAGCCCAATAAGTTTGCTATCTGTTGATCCTCACATTATTTTACTCACAGAAATTCACAATACCTTTATTTTTGTTTGAGTTTGAAGTATACAATTTCTTTAAAATGTAAAATTTGAAATCTCAACAATAAGATATGTTATTGATCCTTGCAATTATGGGTAGATTGCGAATTAAACTATCTTGTCTTTGCTTACAACAGTCATTTTGTTTATAAACTAATTATACATAAATCCTAACTGATAGATAGTTTATAAAGATGAATAATGAACATAGGTCATATATTAAAAAAACAAAAAACAAAAAAAAACTAAACAAGATGAGCGAGTCAAAAATAGTCTTAACAAAAGAATATATATATATGTATATATCATATTTGATTTGTCTATTTTTAATTTTGAAAAAACTAAGTTAATCGATATATAATATGAAGGCATAATGCATAAATATGTCCTTTAACTTGGTTTTAAATCACATTTATACCTCTTCGACTTTGGGTGTATACAAACAAACACTTAAACTTATATAATGTTGAACAAATAGATATATATGTCCTACATGTCATTTTTCGTCCTAAATGGTGTCCTAAGTGTATTGTGTCACGCAGGACTCATGTGTCTATTTGTTCAAATTTATACAAGTTTAAGTGCTTACTTATGTATAAACAAAGTTGAATGACATAAATGTGAAATAAAATCAAATTAAAGGGCATATTTATGCATTATACCTAATACGAAAATCCATATTATTCACTAAAAAATGAGTCGGATTATATGATTACTTTTTTATTCATTTTGCCAATCGTATCCTACGACATTGTTTTTAATTTGCAGACACAATTCATGCTGGATCAGCTTGCAGATCTTCAACAAAAGGTAATTATAAAATTCTACAAATTTCCAATAATTAATAAATGGAATAATTATGCGCGAGAAAT

CTACCCTATGTAGGCGGAATCCTCTTTTCGACTCTGACTCTCCCACTCCAGTCGTGAAAAAACAACAAACTAGTCAAAGGACAGCCTGCCTTATTCTTCTCCCGTTCGGGACCCCTATTTTCTCGGAGATAGCCTGGTCTGAGCTAGAACAGCAGATTCGTGAGCAAGAGCGTATTTCACAGCTGATTCAACAACAGCCATTTTTTCTGGGACCCGCAATTCCGTAGAAAGACATCACGATTCCTTGTGGACGGGGAATCGGCAGAAAGAGATGGGTCGGATACTGGAATCTGCCCAAAAGTCCTGACTTCTATTTAAAATTTACGATGAATTTTAATTTTACAGGAGCAAATGCTTGCAGAATCTAATAGATTACTCCGTAGAAAGGTAAACTAACTTGATAGCCGTGCGTAATGAATAACTTATTTTATTTTCAAAATTATAAATCTAAATACTTAGGTAACTCGATAACATAAGAAGTATTTATACTGATGATATTGGTGTTGTGTTTTTTTTTATTAGTTAGAAGAAAGTGTAGCTGGATTTCCACTTCGATTGTGTTGGGAAGATGGAGGTGATCATCAACTTATGCATCAACAAAATCGTCTCCCTAACACAGAGGGTTTCTTTCAGCCTCTTGGATTGCATTCTTCTTCTCCACATTTTGGGTAATTACTTTTATTATTATTAAAAATAATTTCAATTTTTTTTACTTTTATTTCGATTAATAAATCAATGTGCACCAAGGTACGGTCTAACATAAACAAAAATGTGGGGAATGCTCTTAAAGCCCTAACAAAAGTTATTTGGTACGTGTACTAATGTAATCGTACTATATATCTTACTTGATTAGTGGATGGACAGTACTGGGCACACACAATTGACATAAGTTATTATAAGGAAAAAAAAAGGCCAATAATCAATATAGTCCAACATTACATTATTTATTATAACAGGTCACTCTAGATTAAATGTTAATGAATAACAAAAAGTCTCATATTGATGATTAATGTGATGGGTGGGCTTCTTATAAGGCTTTGACAATCCTACTCTCTTTGAGCTAGTTTTGGGGGTGTGACCTAATTCAACAGAACGTAGTTAAGATTGTGAAGTAAAGTTGATCATTGTTATAACAGGTTTAAATACTTCTAGTAAAAATAGTTCCTAGATAATCCATCGCAAAATAGCTCCTATATAGTTAGTTGGATTTTCATATAATCTATAGCTTATACATAGCTAAATGGGAATAGATGAGAGTTTCTGTTGTTTAGATATGATATTTGATCGGTTTCTAAATCGTTACTATCATGTAGTGAATAATTTTCATGTTATTACTATTACATTTGATTGTTTCTGTGGTTATTTTTTTTTCTAGGTACAATCCTGTTAATACAGATGAGGTGAATGCAGCGGCAACTGCACACAATATGAATGGATTTATTCATGGATGGATGCTTTAAMutant Solyc03g114840 gene allele ej2^(CR) >allele-1 (SEQ ID NO: 15)ATGGGAAGAGGAAGAGTTGAGCTTAAGAGAATAGAAAATAAAATAAATAGGCAAGTCACTTTTGCTAAGAGAAGAAATGGACTTCTTAAAAAAGCTTATGAACTTTCTGTTCTTTGTGATGCTGAAGTTGCCCTTATAATCTTCTCTAATAGGGGTAAACTCTATGAATTTTGCAGCACTTCAAGGTATTTTTTATTTTATTATATTAACATCAAAGATTTTATTTTTTTAAAAAAAACCTTAAGTCCTTCATTACCAAAACCCTTAATTGATTTACAAAGTACTTTCATTAAATTTAGTAATTCTTTTTTTTTTTATCTCTGACTTCAATTATAATGCAAGATCTATGTTGTCTTTATATATATTGAATTATATATGTACTGTATTTTTACTATATACATATAAGATCCTTTTTTCTTTTTTTTCTGTCTCTTTATATAAATATATTTTAAATAGTTGATTTTGAAAGATCTACTAATGTATATTTATTTTTGGAACTTTTGTGTATATGGAATTTTTTTCTTTTTTATGTTTTTTTTTTGTTCTAATTGTTTTAAAAGCGTTTAAGATCAGAATGTTCTTGATTATTCTTTTAGGAAAAAGATTTCCCATACATTGAGTTATTTTTTGATCTGTAGATTGAATTTTTTTAATGAGTTCCGATAGATTTTCGTTCAATTTTTCAATGAAACTATTGAGGGTTGATGATTAGATAATTACTCGATTGAAAGTTTTTATTTCAAAAAAATTATAATTCTTCTTAATTTTATATTTATGAGATAGAGTTAGTTTAGTGATTATATGAAAAATCGTATCAGATTATTGGGAATCGAAACTTAAAAATTCTGAAAATATTATTATAAATTTTACATGTTACAATATTTTTACTGTTAAGATTTGATTTGCAGACTAGGTGTCATGTTTGACAGTTGATAAAAAATCTGTTATTTTTGTTCTTTAATTCCCAAGACGGATAAACAAAGGCTGCTTATGTTGGTTTCCAATAAGCAGCCATAATTTTAAATATTTTTGTTAAGATTAATTAATAACAATTATTTCCACCAGATAATTTTCAAAATTTGTGACCCCGAGTTCATATAAATTGTTAATTTTACTGCTAGAAATTACATCGATAATAATTTATTTAGTGTAATCTTATAAATACGAGGGCAGTAGTGTATAGACTGTTTTTTATTAATCCTGACTCAAAGTGAGGTAAGTTAAGTATATTTTGATTAAAAGGACTACATTTCATTTATGTATGTTTAATTAATATTATTTTGTAAGTCAATAAATCTAAACAACATGAGTTTATCTAGACCCTTAATTATGCACCTTCATTATCAATTTTTTCAATACTCTCCTCAGAACATATGCTTCTCTATAATTTTGTGCACGAGTTAATCAATTCTTCCTTTTCAATAATTAAATATGTGATTTATGTTTAGCACTTATTTTTCGGTTAGTTAATTGATAATAGGAAAAAGCCTCTTTTTTTTTGTGTGTGTGGTAATTAGGATCTTTATTGAATTTAAAATGACCTACTATAGAACTTGGGAGTTTTTCTTCATAATAATGCACTGCAACGTGTTAAAAAAAAAGAATCAAATGAAATTAATAGATGTTTACTGGATTGCCATGGTAAAGTGATAAGTATTAATTTCGCTTTAACTAAGAGATCATTATATTCAAGTCCCCTTGATACAAACTTGCCTTTGTAAATAAGTGTTTTATTTTTCAATGTGAAACTTTCGCTGTTAATTTAAATTTAATTATACTTCTATATAAATACCAAACAATAATGTAATAAAACAAAAAATAAAAGAGTAGATGTTTCATATTGTTAATGCAGCATGGTGAAAACAATTGAAAAGTACCAACGTTGCAGCTATGCTACTTTGGAAGCCAACCAATCAGTTACTGATACTCAGGTACTGCTTTATATTTTAATTTATTTGGCTTTTTTTTAAAAAAATAATTAGTTTTGATTAATATGCATCATTTTATTTATTTTTGGCAACTCTTTATTTATCAGTAATAAGTAATAACTTTTTAACTAGTATATTTAAAAATCACAAAATTTAAGAATATTTTAATAGATTCGACATATTTTAGTTTAAAAATAACAAATTAAATTATGTTTTTAATTTTTTAAATATTCTTACTATAATTATCATGTACTCTTTGATCTGTTCATCTTTTCCATGATAATATTATTTGGTCAGTTAGTGACATAAGAGTTTGAAATTTAGAAAAAAGGAATATTTGGAGAAAACTGAAATGGATATTTAGAAATGAAAGTTATTTAATATAAATATAAGTATGGGCTGCTGAGTTGGGAATCCACGCTGGAGATCTCAAGTTTGAAGCGTCTCACAAACAATAGTAATGTCTTTTTGGTCGAGTTTGTCGGATTGGACTTGTCCGTGGCCTGTGGGTTACTTTTCCTATATGGTTTGCAAGCTATCGGGAATTTTATCCTGGCGCACCCAAATTTGAGTTATTTTTGAGTTTTTATATGAAATAGCTTTGTGAATTCATCGAACTCCCGAAAACATTGAACTTTACTCCAAGTTGAATTGCAGTAAAATAATAGTAGCGATTCTTTAATTTATCCTAACAGTTTTTCGAAATAATAATCCCAAAAAAGTTTAAAATAACCATACCATAAACTTACTGGGTAAGATATTATCTGTCTAATAATATATAGTAGTTTCTTTTGTTTTATTAGTTTATCTAATCCATATTTCATTTCTTGATAAGTTATTCTTAATAGGAAAATAAACTTATTTCGAAAAACTGTTTTTAAAATTTTCTTGAGTTGAGTCTTGGATGAAAAATAGTTAATTTTGCATTAATTAATTTTGTTCTAACAAAAACTAATTAAATTTTTTTGAAGCGCATATTCACTCAAAAAATAAATAAAAACCATCATGCATACAGGAAATGTTCTTTTTTTAATTTATTTTTTCATTGGAGCCCTGACTAATTTTATATCGGTTCATACTTTCATAAATTACAAAAAGTTCAAAATTTAAACTAACCATATAAGTGAATAAAATAAATCAACAAAATATTCACCACATAATACTTTTTAAATAGAATTTTTCATACCAAAGACCTTACTTTAATTAATTAGGGTGAGAGAATCCTATAAGTCAATGCAAAACAATTCTATCTATCGGATTATAATCGTTGATTCATAAAATTTTAAAATCGACGATTTTCATTTAAATGACCCTTTTTTTTCTTTCATTTTTTATTGTTATTCATCTATTTAACTTGTGAGCATCTTTCATATTGATATTTCAGACCCTTAAATTAATTGTTTTCTTACAGAATAACTACCACGAATATCTGAGGCTAAAAGCTAGAGTTGAGCTCCTCCAACGATCTCAGAGGTAATTTCTGTTCACTATCTTTATCTCAAATGAATTCTCATGTTTTTATTTTTCGAGATTCAGATTAAATATAATTTGATGTATTATTAATTTAAATACGTTATTTAATATGGTCCTTATGTCCAACCATTGATTTAATTTGATATTTTTTTAATGAAAATTACACAGAAACTTTCTTGGTGAAGATTTGGGCACCTTGAGCAGCTTGAGAATCAATTAGAGTCTTCCTTAAAGTCAAGGAAGGTAAATTATTTAATCTAATTATACAGAAAAATCATCTAAAAGTTACCTTAATTGCTAGCCCAATAAGTTTGCTATCTGTTGATCCTCACATTATTTTACTCACAGAAATTCACAATACCTTTATTTTTGTTTGAGTTTGAAGTATACAATTTCTTTAAAATGTAAAATTTGAAATCTCAACAATAAGATATGTTATTGATCCTTGCAATTATGGGTAGATTGCGAATTAAACTATCTTGTCTTTGCTTACAACAGTCATTTTGTTTATAAACTAATTATACATAAATCCTAACTGATAGATAGTTTATAAAGATGAATAATGAACATAGGTCATATATTAAAAAAACAAAAAACAAAAAAAAACTAAACAAGATGAGCGAGTCAAAAATAGTCTTAACAAAAGAATATATATATATGTATATATCATATTTGATTTGTCTATTTTTAATTTTGAAAAAACTAAGTTAATCGATATATAATATGAAGGCATAATGCATAAATATGTCCTTTAACTTGGTTTTAAATCACATTTATACCTCTTCGACTTTGGGTGTATACAAACAAACACTTAAACTTATATAATGTTGAACAAATAGATATATATGTCCTACATGTCATTTTTCGTCCTAAATGGTGTCCTAAGTGTATTGTGTCACGCAGGACTCATGTGTCTATTTGTTCAAATTTATACAAGTTTAAGTGCTTACTTATGTATAAACAAAGTTGAATGACATAAATGTGAAATAAAATCAAATTAAAGGGCATATTTATGCATTATACCTAATACGAAAATCCATATTATTCACTAAAAAATGAGTCGGATTATATGATTACTTTTTTATTCATTTTGCCAATCGTATCCTACGACATTGTTTTTAATTTGCAGACACAATTCATGCTGGATCAGCTTGCAGATCTTCAACAAAAGGTAATTATAAAATTCTACAAATTTCCAATAATTAATAAATGGAATAATTATGCGCGAGAAATTTATCTATTTAAAATTTACGATGAATTTTAATTTTACAGGAGCAAATGCTTGCAGAATCTAATAGATTACTCCGTAGAAAGGTAAACTAACTTGATAGCCGTGCGTAATGAATAACTTATTTTATTTTCAAAATTATAAATCTAAATACTTAGGTAACTCGATAACATAAGAAGTATTTATACTGATGATATTGGTGTTGTGTTTTTTTTTATTAGTTAGAAGAAAGTGTAGCTGGATTTCCACTTCGATTGTGTTGGGAAGATGGAGGTGATCATCAACTTATGCATCAACAAAATCGTCTCCCTAACACAGAGGGTTTCTTTCAGCCTCTTGGATTGCATTCTTCTTCTCCACATTTTGGGTAATTACTTTTATTATTATTAAAAATAATTTCAATTTTTTTTACTTTTATTTCGATTAATAAATCAATGTGCACCAAGGTACGGTCTAACATAAACAAAAATGTGGGGAATGCTCTTAAAGCCCTAACAAAAGTTATTTGGTACGTGTACTAATGTAATCGTACTATATATCTTACTTGATTAGTGGATGGACAGTACTGGGCACACACAATTGACATAAGTTATTATAAGGAAAAAAAAAGGCCAATAATCAATATAGTCCAACATTACATTATTTATTATAACAGGTCACTCTAGATTAAATGTTAATGAATAACAAAAAGTCTCATATTGATGATTAATGTGATGGGTGGGCTTCTTATAAGGCTTTGACAATCCTACTCTCTTTGAGCTAGTTTTGGGGGTGTGACCTAATTCAACAGAACGTAGTTAAGATTGTGAAGTAAAGTTGATCATTGTTATAACAGGTTTAAATACTTCTAGTAAAAATAGTTCCTAGATAATCCATCGCAAAATAGCTCCTATATAGTTAGTTGGATTTTCATATAATCTATAGCTTATACATAGCTAAATGGGAATAGATGAGAGTTTCTGTTGTTTAGATATGATATTTGATCGGTTTCTAAATCGTTACTATCATGTAGTGAATAATTTTCATGTTATTACTATTACATTTGATTGTTTCTGTGGTTATTTTTTTTTCTAGGTACAATCCTGTTAATACAGATGAGGTGAATGCAGCGGCAACTGCACACAATATGAATGGATTTATTCATGGATGGATGCTTTAA >allele-3 (SEQ ID NO: 16)ATGGGAAGAGGAAGAGTTGAGCTTAAGAGAATAGAAAATAAAATAAATAGGCAAGTCACTTTTGCTAAGAGAAGAAATGGACTTCTTAAAAAAGCTTATGAACTTTCTGTTCTTTGTGATGCTGAAGTTGCCCTTATAATCTTCTCTAATAGGGGTAAACTCTATGAATTTTGCAGCACTTCAAGGTATTTTTTATTTTATTATATTAACATCAAAGATTTTATTTTTTTAAAAAAAACCTTAAGTCCTTCATTACCAAAACCCTTAATTGATTTACAAAGTACTTTCATTAAATTTAGTAATTCTTTTTTTTTTTATCTCTGACTTCAATTATAATGCAAGATCTATGTTGTCTTTATATATATTGAATTATATATGTACTGTATTTTTACTATATACATATAAGATCCTTTTTTCTTTTTTTTCTGTCTCTTTATATAAATATATTTTAAATAGTTGATTTTGAAAGATCTACTAATGTATATTTATTTTTGGAACTTTTGTGTATATGGAATTTTTTTCTTTTTTATGTTTTTTTTTTGTTCTAATTGTTTTAAAAGCGTTTAAGATCAGAATGTTCTTGATTATTCTTTTAGGAAAAAGATTTCCCATACATTGAGTTATTTTTTGATCTGTAGATTGAATTTTTTTAATGAGTTCCGATAGATTTTCGTTCAATTTTTCAATGAAACTATTGAGGGTTGATGATTAGATAATTACTCGATTGAAAGTTTTTATTTCAAAAAAATTATAATTCTTCTTAATTTTATATTTATGAGATAGAGTTAGTTTAGTGATTATATGAAAAATCGTATCAGATTATTGGGAATCGAAACTTAAAAATTCTGAAAATATTATTATAAATTTTACATGTTACAATATTTTTACTGTTAAGATTTGATTTGCAGACTAGGTGTCATGTTTGACAGTTGATAAAAAATCTGTTATTTTTGTTCTTTAATTCCCAAGACGGATAAACAAAGGCTGCTTATGTTGGTTTCCAATAAGCAGCCATAATTTTAAATATTTTTGTTAAGATTAATTAATAACAATTATTTCCACCAGATAATTTTCAAAATTTGTGACCCCGAGTTCATATAAATTGTTAATTTTACTGCTAGAAATTACATCGATAATAATTTATTTAGTGTAATCTTATAAATACGAGGGCAGTAGTGTATAGACTGTTTTTTATTAATCCTGACTCAAAGTGAGGTAAGTTAAGTATATTTTGATTAAAAGGACTACATTTCATTTATGTATGTTTAATTAATATTATTTTGTAAGTCAATAAATCTAAACAACATGAGTTTATCTAGACCCTTAATTATGCACCTTCATTATCAATTTTTTCAATACTCTCCTCAGAACATATGCTTCTCTATAATTTTGTGCACGAGTTAATCAATTCTTCCTTTTCAATAATTAAATATGTGATTTATGTTTAGCACTTATTTTTCGGTTAGTTAATTGATAATAGGAAAAAGCCTCTTTTTTTTTGTGTGTGTGGTAATTAGGATCTTTATTGAATTTAAAATGACCTACTATAGAACTTGGGAGTTTTTCTTCATAATAATGCACTGCAACGTGTTAAAAAAAAAGAATCAAATGAAATTAATAGATGTTTACTGGATTGCCATGGTAAAGTGATAAGTATTAATTTCGCTTTAACTAAGAGATCATTATATTCAAGTCCCCTTGATACAAACTTGCCTTTGTAAATAAGTGTTTTATTTTTCAATGTGAAACTTTCGCTGTTAATTTAAATTTAATTATACTTCTATATAAATACCAAACAATAATGTAATAAAACAAAAAATAAAAGAGTAGATGTTTCATATTGTTAATGCAGCATGGTGAAAACAATTGAAAAGTACCAACGTTGCAGCTATGCTACTTTGGAAGCCAACCAATCAGTTACTGATACTCAGGTACTGCTTTATATTTTAATTTATTTGGCTTTTTTTTAAAAAAATAATTAGTTTTGATTAATATGCATCATTTTATTTATTTTTGGCAACTCTTTATTTATCAGTAATAAGTAATAACTTTTTAACTAGTATATTTAAAAATCACAAAATTTAAGAATATTTTAATAGATTCGACATATTTTAGTTTAAAAATAACAAATTAAATTATGTTTTTAATTTTTTAAATATTCTTACTATAATTATCATGTACTCTTTGATCTGTTCATCTTTTCCATGATAATATTATTTGGTCAGTTAGTGACATAAGAGTTTGAAATTTAGAAAAAAGGAATATTTGGAGAAAACTGAAATGGATATTTAGAAATGAAAGTTATTTAATATAAATATAAGTATGGGCTGCTGAGTTGGGAATCCACGCTGGAGATCTCAAGTTTGAAGCGTCTCACAAACAATAGTAATGTCTTTTTGGTCGAGTTTGTCGGATTGGACTTGTCCGTGGCCTGTGGGTTACTTTTCCTATATGGTTTGCAAGCTATCGGGAATTTTATCCTGGCGCACCCAAATTTGAGTTATTTTTGAGTTTTTATATGAAATAGCTTTGTGAATTCATCGAACTCCCGAAAACATTGAACTTTACTCCAAGTTGAATTGCAGTAAAATAATAGTAGCGATTCTTTAATTTATCCTAACAGTTTTTCGAAATAATAATCCCAAAAAAGTTTAAAATAACCATACCATAAACTTACTGGGTAAGATATTATCTGTCTAATAATATATAGTAGTTTCTTTTGTTTTATTAGTTTATCTAATCCATATTTCATTTCTTGATAAGTTATTCTTAATAGGAAAATAAACTTATTTCGAAAAACTGTTTTTAAAATTTTCTTGAGTTGAGTCTTGGATGAAAAATAGTTAATTTTGCATTAATTAATTTTGTTCTAACAAAAACTAATTAAATTTTTTTGAAGCGCATATTCACTCAAAAAATAAATAAAAACCATCATGCATACAGGAAATGTTCTTTTTTTAATTTATTTTTTCATTGGAGCCCTGACTAATTTTATATCGGTTCATACTTTCATAAATTACAAAAAGTTCAAAATTTAAACTAACCATATAAGTGAATAAAATAAATCAACAAAATATTCACCACATAATACTTTTTAAATAGAATTTTTCATACCAAAGACCTTACTTTAATTAATTAGGGTGAGAGAATCCTATAAGTCAATGCAAAACAATTCTATCTATCGGATTATAATCGTTGATTCATAAAATTTTAAAATCGACGATTTTCATTTAAATGACCCTTTTTTTTCTTTCATTTTTTATTGTTATTCATCTATTTAACTTGTGAGCATCTTTCATATTGATATTTCAGACCCTTAAATTAATTGTTTTCTTACAGAATAACTACCACGAATATCTGAGGCTAAAAGCTAGAGTTGAGCTCCTCCAACGATCTCAGAGGTAATTTCTGTTCACTATCTTTATCTCAAATGAATTCTCATGTTTTTATTTTTCGAGATTCAGATTAAATATAATTTGATGTATTATTAATTTAAATACGTTATTTAATATGGTCCTTATGTCCAACCATTGATTTAATTTGATATTTTTTTAATGAAAATTACACAGAAACTTTCTTGGTGAAGATTTGGGCACGTTAAGCTTCGAAGGACCTTGAGCAGCTTGAGAATCAATTAGAGTCTTCCTTAAAGCAAATCAGGTCAAGGAAGGTAAATTATTTAATCTAATTATACAGAAAAATCATCTAAAAGTTACCTTAATTGCTAGCCCAATAAGTTTGCTATCTGTTGATCCTCACATTATTTTACTCACAGAAATTCACAATACCTTTATTTTTGTTTGAGTTTGAAGTATACAATTTCTTTAAAATGTAAAATTTGAAATCTCAACAATAAGATATGTTATTGATCCTTGCAATTATGGGTAGATTGCGAATTAAACTATCTTGTCTTTGCTTACAACAGTCATTTTGTTTATAAACTAATTATACATAAATCCTAACTGATAGATAGTTTATAAAGATGAATAATGAACATAGGTCATATATTAAAAAAACAAAAAACAAAAAAAAACTAAACAAGATGAGCGAGTCAAAAATAGTCTTAACAAAAGAATATATATATATGTATATATCATATTTGATTTGTCTATTTTTAATTTTGAAAAAACTAAGTTAATCGATATATAATATGAAGGCATAATGCATAAATATGTCCTTTAACTTGGTTTTAAATCACATTTATACCTCTTCGACTTTGGGTGTATACAAACAAACACTTAAACTTATATAATGTTGAACAAATAGATATATATGTCCTACATGTCATTTTTCGTCCTAAATGGTGTCCTAAGTGTATTGTGTCACGCAGGACTCATGTGTCTATTTGTTCAAATTTATACAAGTTTAAGTGCTTACTTATGTATAAACAAAGTTGAATGACATAAATGTGAAATAAAATCAAATTAAAGGGCATATTTATGCATTATACCTAATACGAAAATCCATATTATTCACTAAAAAATGAGTCGGATTATATGATTACTTTTTTATTCATTTTGCCAATCGTATCCTACGACATTGTTTTTAATTTGCAGACACAATTCATGCTGGATCAGCTTGCAGATCTTCAACAAAAGGTAATTATAAAATTCTACAAATTTCCAATAATTAATAAATGGAATAATTATGCGCGAGAAATTTATCTATTTAAAATTTACGATGAATTTTAATTTTACAGGAGCAAATGCTTGCAGAATCTAATAGATTACTCCGTAGAAAGGTAAACTAACTTGATAGCCGTGCGTAATGAATAACTTATTTTATTTTCAAAATTATAAATCTAAATACTTAGGTAACTCGATAACATAAGAAGTATTTATACTGATGATATTGGTGTTGTGTTTTTTTTTATTAGTTAGAAGAAAGTGTAGCTGGATTTCCACTTCGATTGTGTTGGGAAGATGGAGGTGATCATCAACTTATGCATCAACAAAATCGTCTCCCTAACACAGAGGGTTTCTTTCAGCCTCTTGGATTGCATTCTTCTTCTCCACATTTTGGGTAATTACTTTTATTATTATTAAAAATAATTTCAATTTTTTTTACTTTTATTTCGATTAATAAATCAATGTGCACCAAGGTACGGTCTAACATAAACAAAAATGTGGGGAATGCTCTTAAAGCCCTAACAAAAGTTATTTGGTACGTGTACTAATGTAATCGTACTATATATCTTACTTGATTAGTGGATGGACAGTACTGGGCACACACAATTGACATAAGTTATTATAAGGAAAAAAAAAGGCCAATAATCAATATAGTCCAACATTACATTATTTATTATAACAGGTCACTCTAGATTAAATGTTAATGAATAACAAAAAGTCTCATATTGATGATTAATGTGATGGGTGGGCTTCTTATAAGGCTTTGACAATCCTACTCTCTTTGAGCTAGTTTTGGGGGTGTGACCTAATTCAACAGAACGTAGTTAAGATTGTGAAGTAAAGTTGATCATTGTTATAACAGGTTTAAATACTTCTAGTAAAAATAGTTCCTAGATAATCCATCGCAAAATAGCTCCTATATAGTTAGTTGGATTTTCATATAATCTATAGCTTATACATAGCTAAATGGGAATAGATGAGAGTTTCTGTTGTTTAGATATGATATTTGATCGGTTTCTAAATCGTTACTATCATGTAGTGAATAATTTTCATGTTATTACTATTACATTTGATTGTTTCTGTGGTTATTTTTTTTTCTAGGTACAATCCTGTTAATACAGATGAGGTGAATGCAGCGGCAACTGCACACAATATGAATGGATTTATTCATGGATGGATGCTTTAA

EXAMPLES Example 1. Bypassing Negative Epistasis on Yield in TomatoImposed by a Domestication Gene Abstract

Selection for inflorescence architecture with improved flower productionand yield is common to many domesticated crops. However, tomatoinflorescences resemble wild ancestors, and breeders avoided excessivebranching because of low fertility. The present disclosure relates tothe finding of branched variants that carry mutations in two relatedtranscription factors that had been selected independently. As describedherein, one founder mutation enlarged the leaf-like organs on fruits andwas selected as fruit size increased during domestication. The othermutation eliminated the flower abscission zone, providing “jointless”fruit stems that reduced fruit dropping and facilitated mechanicalharvesting. Stacking both beneficial traits caused undesirable branchingand sterility due to epistasis, which breeders overcame withsuppressors. However, this restricted the opportunity for productivitygains from weak branching. Exploiting natural and engineered alleles formultiple family members, we achieved a continuum of inflorescencecomplexity that allowed breeding of higher yielding hybrids.Characterizing and neutralizing similar cases of negative epistasiscould improve productivity in many agricultural organisms.

Methods Experimental Model and Subject Details Plant Material and GrowthConditions

Seeds of the standard S. lycopersicum cultivar M82 (LA3475) were fromthe present stocks. Core collection germplasm (www.eu-sol.wur.nl) wasfrom the seed stocks of Z. Lippman, D. Zamir, and S. Huang (Lin et al.,2014). Seeds of the jointless S. cheesmaniae accession LA0166 wereobtained from the Charles M. Rick Tomato Genetics Resource Center (TGRC)at the University of California, Davis. The frondea mutant was obtainedfrom the gene bank of the Leibniz Institute of Plant Genetics and CropPlant Research (IPK) in Gatersleben, Germany. Seed of the longinflorescence (lin) mutant in the Micro-tom background (TOM-JPG5091) wasprovided by the University of Tsukuba, Gene Research Center, through theNational Bio-Resource Project (NBRP) of the AMED, Japan(tomatoma.nbrp.jp/). The lin mutant was backcrossed four times to thestandard M82 cultivar. The landrace collection (S. lycopersicunm var.cerasifornme) was from the seed stocks of E. van der Knaap. Tissuesamples, DNA, or seed of elite breeding lines were obtained fromSyngenta, Nunhems, Monsanto, Lipman Seeds, Johnny's Seeds, andTomatoGrowers.

Seeds were either pre-germinated on moistened Whatman paper at 28° C. incomplete darkness or directly sown and germinated in soil in 96-cellplastic flats. Plants were grown under long-day conditions (16-hlight/8-h dark) in a greenhouse under natural light supplemented withartificial light from high-pressure sodium bulbs (˜250 μmol m⁻² s⁻¹).Daytime and nighttime temperatures were 26-28° C. and 18-20° C.,respectively, with a relative humidity of 40-60%.

Analyses of inflorescence architecture, sepal length, fruit type, andproductivity traits were conducted on plants grown in the fields at ColdSpring Harbor Laboratory, Cold Spring Harbor, N.Y., the Cornell LongIsland Horticultural Experiment Station in Riverhead, N.Y., and nethouses in Hatzav, Israel. Analyses of sepal length in the landraces wereconducted on plants grown in the fields of the Durham horticulture farmat the University of Georgia, Athens, Ga. Seeds were germinated in96-cell flats and grown for 32 d in the greenhouse before beingtransplanted to the field. Plants were grown under drip irrigation andstandard fertilizer regimes. Damaged or diseased plants were markedthroughout the season and were excluded from the analyses.

Method Details Plant Phenotyping

For analyses of sepal length, the length of sepals and petals of 10closed flower buds per accession were manually measured and thesepal/petal ratio was calculated. Mature floral buds of similardevelopmental stage were collected (1-2 days before anthesis, i.e.before flower opening). For analyses of inflorescence complexity, thenumber of branching events was counted on at least 5 inflorescences oneach replicate plant.

Yeast Two-Hybrid Analysis

Protein interaction assays in yeast were performed using the MatchmakerGold Yeast Two-Hybrid System (Clontech) as described before (Park etal., 2014b). The coding sequences for bait proteins were cloned into thepGBKT7 vector, and the resulting vectors were transformed into theY2HGold yeast strain. The coding sequences for prey proteins were clonedinto the pGADT7 AD vector, and the resulting vectors were transformedinto the Y187 yeast strain. After mating the two yeast strainsexpressing bait and prey proteins, diploid yeast cells were selected andgrown on dropout medium without leucine and tryptophan. To assayprotein-protein interactions, clones were grown on triple-dropout mediumwithout leucine, tryptophan, and histidine for 3 d at 30° C. To blockauto-activation, 3 mM 3-amino-1,2,4-triazole (3-AT) were added oradenine was removed from the triple-dropout medium. All primer sequencesused for cloning can be found in Table 2.

Meristem Imaging

Live meristems were imaged using a Nikon SMZ1500 stereomicroscope(Nikon). Shoot apices were dissected from seedlings and older leafprimordia were removed to expose meristems. Immediately afterdissection, sequences of optical layers were imaged using a Nikon DS-Ri1digital camera (Nikon) mounted on the stereomicroscope. Z-stacks ofoptical sections were aligned and merged to produce final focused imagesusing the NIS Elements BR3.2 software (Nikon).

Meristem Transcriptome Profiling

Meristem collection, RNA extraction, and library preparation for s2mutant plants was performed as previously described (Park et al., 2012).Briefly, seedling shoots were collected at the vegetative meristem (VM),transition meristem (TM), sympodial inflorescence meristem (SIM), andfloral meristem (FM) stage of meristem maturation, and immediately fixedthem in ice-cold acetone. Meristems were manually dissected under astereoscope and two biological replicates consisting of 30-50 meristemsfrom independent plants were generated. Total RNA was extracted with thePicoPure RNA Extraction kit (Arcturus) and mRNA was purified withDynabeads mRNA Purification kits (Thermo Fisher). Barcoded librarieswere prepared using the NEBNext Ultra RNA library prep kit for Illuminaaccording to the manufacturer's instructions, and assessed for sizedistribution and concentration with a Bioanalyzer 2100 (Agilent) and theKapa Library quantification kit (Kapa Biosystems), respectively.Libraries were sequenced on a single Illumina Hiseq 2500 lane(222,279,510 million paired-end reads) at the Genome Center of ColdSpring Harbor Laboratories, Cold Spring Harbor.

Previously collected reads for wild-type tomato cultivar M82, compoundinflorescence (s) mutant (Lemmon et al., 2016; Park et al., 2012), andreads for the s2 mutant were trimmed by quality using Trimmomatic(Bolger et al., 2014b) and aligned to the reference genome sequence oftomato (SL2.50) (Consortium, 2012) using Tophat2 (Kim et al., 2013).Alignments were sorted with samtools (Li et al., 2009) and geneexpression quantified as unique read pairs aligned to referenceannotated gene features (ITAG2.4) using HTSeq-count (Anders et al.,2015).

All statistical analyses of gene expression were conducted in R (RTeam,2015). Expression of individual genes is shown as transcripts permillion (TPM). Significant differential expression between meristemstages in wild-type tomato cultivar M82 was identified for 2,582 geneswith edgeR (Robinson et al., 2009) using 2-foldchange, average 1 CPM,and FDR≤0.10 cutoffs (Lemmon et al., 2016). To compare expressiondynamics by principal component analysis (PCA), z-score normalization ofraw counts was used within genotype to minimize the impact of thedifferent sequencing lengths (50 bp vs. 100 bp) and platforms (GAIIx andHiSeq2500). PCA was conducted on normalized expression values for the2,582 dynamic genes in wild-type tomato cultivar M82, s, and s2 usingthe prcomp function in R (RTeam, 2015). The first two principalcomponents were then plotted to assess acceleration or delay of themeristem maturation process in mutant samples. The proportion of TM andFM marker genes with moderate and severely delayed expression wasassessed by a two-step k-means clustering. First, normalized WTexpression was grouped into twelve clusters and the two clusters withthe most specific TM and FM expression were designated as markers.Mutant expression from TM and FM marker genes was normalized with WT,producing WT:s and WT:s2 normalized expression datasets. Finally,k-means clustering (12 clusters) was performed on s and s2 normalizedexpression alone and clusters with delays in activation compared to WTwere identified by hand.

Mapping-by-Sequencing

To map the causal mutations in the s2 mutant, two second-generation (F₂)populations were generated by crossing s2 with the S. lycopersicumcultivar M82, and s2 with S. pimpinellifolium. From a total of 464s2×M82 F₂ plants, 25 s2 mutants, 20 j2 mutants, and 13 WT siblings wereselected for tissue collection, nuclei isolation, and DNA extraction. Anequal amount of tissue from each plant (˜0.2 g) was pooled for DNAextraction using standard protocols. Libraries were prepared with theIllumina TruSeq DNA PCR-free prep kit from 2 μg genomic DNA sheared to550 bp insert size. From a total of 576 s2×S. pimpinellifolium F₂plants, 16 s2 mutants, 9 j2 mutants, and 13 wild-type siblings wereselected for DNA extraction. DNA was also extracted from the s2 parent(LA4371). Libraries were prepared with the Illumina TruSeq Nano DNA prepkit from 200 ng genomic DNA sheared to 550 bp insert size and 8 cyclesof final amplification. All DNA libraries were sequenced on an IlluminaNextSeq platform at the Cold Spring Harbor Laboratory Genome Center(Woodbury, N.Y.). For the s2×M82 F₂ population, 62,317,992, 73,496,741,and 79,699,274 paired-end 151-bp reads were obtained for the s2 mutant,j2 mutant, and the WT sibling samples, respectively. For the s2×S.pimpinellifolium F₂ population, 32,979,728, 82,439,796, and 50,763,441paired-end 151-bp reads were obtained for pools of s2,j2, and the WTsiblings, respectively. For the s2 parent 48,281,689 paired-end 151-bpreads were obtained.

To map the causal mutation in the in mutant, a F₂ population wasgenerated by crossing the in mutant with S. pimpinellifolium. From atotal of 216 F₂ plants, 8 lin mutant plants were selected with the moststrongly branched inflorescences and 17 WT siblings for tissuecollection. An equal amount of tissue from each plant (˜0.2 g) waspooled for nuclei isolation and DNA extraction using standard protocols.Barcoded libraries were prepared with the Illumina TruSeq DNA PCR-freeprep kit from 2 μg genomic DNA sheared to 550 bp insert size andsequenced as above. 4,624,816 and 5,063,861 paired-end 101-bp reads wereobtained for the in mutant and the WT sibling pools, respectively. Tofind the in mutation, a pool of 7 lin×S. pimpinellifolium F₂ mutantplants was resequenced on the Illumina HiSeq2500 platform, and anadditional 161,827,433 paired-end 101-bp reads were obtained.

To map s2 suppressor loci in S. pimpinellifolium, 1,536 S.pimpinellifolium×s2 F₂ plants were regrown and 92 homozygous j2^(TE) ej2double mutants were selected by PCR genotyping. Primers are listed inTable 2. 18 s2 mutants, 6 moderately suppressed s2 mutants, and 2strongly suppressed s2 mutants were selected for tissue collection,nuclei isolation, and DNA extraction. Libraries were prepared with theIllumina TruSeq DNA PCR-free prep kit from 2 μg genomic DNA sheared to550 bp insert size, and sequenced as above. 38,060,212, 38,044,727 and52,426,078 paired-end 151-bp reads were obtained for the pools of s2,moderately suppressed s2, and the strongly suppressed s2 plants,respectively.

Genomic DNA reads were trimmed by quality using Trimmomatic and pairedreads mapped to the reference tomato genome (SL2.50) using BWA-MEM (Li,2013; Li and Durbin, 2009). Alignments were then sorted with samtoolsand duplicates marked with PicardTools (Li et al., 2009,broadinstitute.github.io/picard). SNPs were called withsamtools/bcftools (Li, 2011; Li et al., 2009) using read alignments forthe various genomic DNA sequencing pools from this project in additionto reference M82 (Bolger et al., 2014a) and S. pimpinellifolium(Consortium, 2012) reads. Called SNPs were then filtered for bi-allelichigh quality SNPs at least 100 bp from a called indel using bcftools(Li, 2011). Following read alignment and SNP calling, all statistics andcalculations were done in R (RTeam, 2015). Read depth for each allele atsegregating bi-allelic SNPs in 1 Mb sliding windows (by 100 kb) wassummed for the various mutant (s2,j2^(TE) or suppression of s2) andwild-type sequencing pools and mutant:non-mutant SNP ratios werecalculated. Finally, mutant SNP ratio was divided by wild-type SNP ratio(+0.5) and plotted across the 12 tomato chromosomes.

Tissue Collection and RNA Extraction

For semi-quantitative RT-PCR, seeds were germinated on moistened Whatmanpaper at 28° C. in complete darkness. Seedlings at similar germinationstages were transferred to soil in 72-cell plastic flats and grown inthe greenhouse. Shoot apices were collected at the floral meristem (FM)stage of meristem maturation (Park et al., 2012), and immediatelyflash-frozen in liquid nitrogen. Total RNA was extracted using theRNeasy Plant Mini Kit (Qiagen) and treated with the RNase Free DNase Set(Qiagen), or the Arcturus PicoPure RNA Extraction kit (Thermo Fisher)according to the manufacturer's instructions. 100 ng to 1 μg of totalRNA was used for cDNA synthesis using the SuperScript III First-StrandSynthesis System (Invitrogen). All primer sequences can be found inTable 2.

Phylogenetic Analyses and Sequence Analyses

Sequences of tomato and Arabidopsis SEP family members were obtainedfrom the Phytozome v11 database (phytozome.net) and aligned using theClustalW function in MEGA. Phylogenetic trees for proteins with 1,000bootstrap replicates were constructed using the maximum likelihoodmethod in MEGA6 (Tamura et al., 2013). Homologous proteins in the cladescontaining Arabidopsis SEP1/2, SEP3, and SEP4 were assigned as SEP1/2-,SEP3-, and SEP4-homologs, respectively.

For analysing linkage between EJ2 and FW3.2, the M9 SNP was genotyped atposition SL2.50ch03:64799226 (Chakrabarti et al., 2013) (G in S.pimpinellifolium (FW3.2) and A in S. lycopersicum cv. M82 (fw3.2)) inaccessions of the tomato core collection using published genomesequencing data (Lin et al., 2014; Tieman et al., 2017).

CRISPR/Cas9 Mutagenesis, Plant Transformation, and Selection of MutantAlleles

CRISPR/Cas9 mutagenesis and generation of transgenic plants wasperformed following the standard protocol (Belhaj et al., 2013; Brookset al., 2014). Briefly, two single-guide (sg)RNAs binding in the codingsequence of the target gene were designed using the CRISPR-P tool(cbi.hzau.edu.cn/cgi-bin/CRISPR) (Lei et al., 2014). Vectors wereassembled using the Golden Gate cloning system (Werner et al., 2012).The sgRNA-1 and sgRNA-2 were cloned downstream of the Arabidopsis U6promoter in the Level 1 acceptors pICH47751 and pICH47761, respectively.The Level1 constructs pICH47731-NOSpro::NPTII, pICH47742-35S:Cas9,pICH47751-AtU6pro:sgRNA-1, and pICH47761-AtU6::sgRNA-2 were assembled inthe binary Level 2 vector pAGM4723. Fifteen-μl restriction-ligationreactions were performed in a thermocycler (3 min at 37° C. and 4 min at16° for 20 cycles, 5 min at 50° C., 5 min at 80° C., and final storageat 4° C.). All sgRNA sequences are listed in Table 2.

Final binary vectors were transformed into the tomato cultivar M82 andthe tomato wild species S. pimpinelifolium by Agrobacteriumtumefaciens-mediated transformation (Gupta, S. and Van Eck, 2016). Afterin-vitro regeneration, culture medium was washed from the root systemand plants transplanted into soil. For acclimation, plants were coveredwith transparent plastic domes and maintained in a shaded area for 5days. A total of 8 first-generation (To) transgenics were genotyped forinduced lesions using forward and reverse primer flanking the sgRNAtarget sites. PCR products were separated on agarose gels and selectedproducts were cloned into pSC-A-amp/kan vector (StrataClone Blunt PCRCloning Kit, Stratagene). At least 6 clones per PCR product weresequenced using M13-F and M13-R primer. T₀ plants with lesions werebackcrossed to wild type and the F₁ generation was genotyped fordesirable large deletion alleles and presence/absence of the CRISPR/Cas9transgene using primer binding the 3′ of the 35S promoter and the 5′ ofthe Cas9 transgene, respectively. All primers are listed in Table 2.Plants heterozygous for the engineered deletion alleles and lacking thetransgene were self-pollinated to isolate homozygous, non-transgenicnull mutants from the F₂ generation.

Generation of Parental and Hybrid Lines for Cherry Tomato Breeding andYield Trials Under Agricultural Greenhouse Conditions

To test the potential of j2 ej2 and s genotypes for fresh-market tomatobreeding, hybrids were generated by crossing near-isogenic linesisolated from a breeding population that was developed for breedinghigh-yielding, indeterminate cherry tomato cultivars with a range offruit shapes (Dani Zamir). Depending on genotype, near-isogenic lineswere generated by backcrossing once to the respective cherry parents(BC₁) followed by inbreeding for 3 generations (F₃) or by inbreeding for3-6 generations (F₃-F₆). Fruit shapes, inflorescence types, and yieldcharacteristics were evaluated and selected each generation. Tenreplicate plants per parental and hybrid line were grown in a randomizedplot design in net houses in Hatzav, Israel in the year 2017. Damaged ordiseased plants were marked throughout the season and were excluded fromthe analyses.

j2 Ej2 Hybrid Experiment

A jointless (j2^(TE)) processing inbred (F₆) wild type for EJ2 (j2 EJ2)served as parent (P-6022) for generating test and control hybrids. Testparents were isolated from a jointless (j2^(TE)) cherry inbredpopulation (BC₁F3), which segregated for ej2^(w). Twoj2^(TE) parents(P-6086-2 and P-6086-9) and two j2^(TE) ej2 parents (P-6086-4 andP-6086-8) were selected by ej2^(w) genotyping, and were crossed toP-6022. Control hybrids were generated by crossing the j2^(TE) testparents (P-6086-2 for trail-I and P-6086-9 for trial-2) to the j2^(TE)parent (P-6022). Test hybrids were generated by bulk crossing thej2^(TE) ej2^(w) test parents (P-6086-4 for trail-1 and P-6086-8 fortrial-2) to the j2^(TE) parent (P-6022).

s Hybrid Experiment

An indeterminate cocktail inbred (F₃) and a determinate cherry inbred(F₃) served as parents (P-6097 and P-6105, respectively) for generatingtest and control hybrids. Test parents were isolated from anindeterminate cherry-type F₅ inbred line that segregated the s mutation.One parent wild type for S (P-6089) and one s mutant parent (P-6090)were selected by phenotyping and self-fertilized. The F₆ generation wasstable for unbranched (P-6089) and compound inflorescences (P-6090).Control and test hybrids were generated by bulk crossing the S parents(P-6097 for trail-1 and P-6105 for trial-2) to the S (P-6089) and the s(P-6090) test parents, respectively.

For analyses of yield component traits, mature green (MG) and red fruits(MR) were collected from 6 subsequent individual inflorescences and MGfruit number (MGFN), MR fruit number (MRFN), MG fruit weight (MGFW), andMR fruit weight (MRFW) was determined per inflorescence. Total fruitnumber (TFN) was the sum of MGFN and MRFN from each plant. Total yield(TY) was the sum of MGFW and MRFW from each plant. The average fruitweight (FW) was calculated by dividing MRFW by MRFN. From each plant,7-10 fruits from at least one inflorescence were randomly selected todetermine total soluble sugar content (Brix) in fruit juice. Brix value(percent) was quantified with a digital Brix refractometer (ATAGOPalette). For each measured yield parameter, mean values and percentagedifference to the control hybrid were statistically compared using twotailed, two-sample t-tests.

Quantification and Statistical Analyses Sampling

For quantitative analyses of flower number per inflorescence andinflorescence internode length, at least 10 inflorescences were analyzedper genotype. For quantitative analyses of inflorescence complexity atleast 5 inflorescences each from 6 individual replicate plants wereanalyzed per genotype. For quantitative analyses of relative sepallength, at least flowers were analyzed per genotype or ecotype. Hybridinflorescence traits (number of branching events per inflorescence,total number of branches and flowers per plant) were determined for 6subsequent inflorescences per individual plant and 9-10 individualplants per hybrid line. Total number of mature green and red fruits perindividual plant was determined from 6 subsequent inflorescences perplant and 9-10 individual plants per hybrid line. Exact numbers ofindividuals (N) are presented in all FIGs. Statistical calculations wereperformed using R and Microsoft Excel. Mean values for each measuredparameter were compared using two-tailed, two-samples Student's t-test.

Transcriptome Quantification

Reads for the wild-type M82, compound inflorescence (s) mutant (Lemmonet al., 2016; Park et al., 2012), and the s2 mutant were trimmed byquality using Trimmomatic v0.32 (HiSeq2500 parameters:ILLUMINACLIP:TruSeq3-PE-2.fa:2:40:15:1:FALSE LEADING:3 TRAILING:3SLIDINGWINDOW:4:15 MINLEN:36; GAIIx parameters:ILLUMINACLIP:TruSeq2-PE.fa:2:30:10:1:FALSE LEADING:3 TRAILING:3SLIDINGWINDOW:4:15 MINLEN:36 TOPHRED33) (Bolger et al., 2014b) andaligned to the reference genome sequence of tomato (SL2.50) (Consortium,2012) using Tophat2 v2.0.127 (parameters:—b2-very-sensitive—read-mismatches 2—read-edit-dist 2—min-anchor8—splice-mismatches 0—min-intron-length 50—max-intron-length50000—max-multihits 20) (Kim et al., 2013). Alignments were sorted withsamtools (Li et al., 2009) and gene expression quantified as unique readpairs aligned to reference annotated gene features (ITAG2.4) usingHTSeq-count v0.6.08 (parameters:—format=bam—order=name—stranded=no—type=exon—idattr=Parent) (Anders etal., 2015).

All statistical analyses of gene expression were conducted in R (RTeam,2015). Significant differential expression between meristem stages inwild-type M82 was identified for 2,582 genes with edgeR (Robinson etal., 2009) using 2-foldchange, average 1 CPM, and FDR≤0.10 cutoffs(Lemmon et al., 2016). To compare expression dynamics between genotypes,z-score normalization was used within genotype to minimize the impact ofthe different sequencing lengths (50 bp vs. 100 bp) and platforms (GAIIxand HiSeq2500). A principal component analysis (PCA) was conducted onthese normalized expression values for the 2,582 dynamic genes includingwild-type M82, s, and s2 using the prcomp function in R (RTeam, 2015).The first two principal components were then plotted to assess modifiedmaturation schedules in the mutant samples. The proportion of TM and FMmarker genes with moderate and strongly delayed expression was assessedby a two-step k-means clustering. First, WT expression (TPM) was z-scorenormalized and clustered into twelve groups using the kmeans2 functionfrom the Mfuzz package (Futschik, 2015) in R. The two clusters with themost specific TM and FM expression (clusters 06 and 08, respectively;FIG. 8A) were designated as marker clusters. Mutant s and s2 expression(TPM) from the 277 TM and 241 FM marker genes was z-score normalizedwith WT expression, producing a WT:s normalized expression and WT:s2normalized expression dataset. Finally, k-means clustering (12 clusters)was performed on s (FIG. 8B) and s2 (FIG. 8C) expression alone(normalized by WT expression levels) and clusters with moderate andsevere delays in activation compared to WT were manually identified.

Mapping

For mapping-by-sequencing of the various mutants, reads were trimmed byquality using Trimmomatic v0.32 (HiSeq 2500 read parameters:ILLUMINACLIP:TruSeq3-PE-2.fa:2:40:15:1:FALSE LEADING:3 TRAILING:3SLIDINGWINDOW:4:15 MINLEN:36; GAIIx read parameters:ILLUMINACLIP:TruSeq2-PE.fa:2:30:10:1:FALSE LEADING:3 TRAILING:3SLIDINGWINDOW:4:15 MINLEN:36 TOPHRED33) and paired reads mapped to thereference tomato genome (SL2.50) using BWA-MEM v0.7.10-r789 (parameters:-M) (Li, 2013). Alignments were then sorted with samtools and duplicatesmarked with PicardTools vi.126 (parameters:VALIDATION_STRINGENCY=LENIENT) (Li et al., 2009,broadinstitute.github.io/picard). SNPs were called withsamtools/bcftools v1.3.1 (samtools mpileup parameters:—ignore-RG—max-depth 1000000—output-tags DP,AN—min-BQ0—no-BAQ—uncompressed—BCF; bcftools call parameters:—multiallelic-caller—variants-only—output-type z) (Li, 2011; Li et al.,2009) using read alignments for the various sequencing pools from thisproject in addition to reference M82 (Bolger et al., 2014a) and S.pimpinellifolium (Consortium, 2012) reads. Called SNPs were thenfiltered for bi-allelic high quality (MQ>=50) SNPs at least 100 bp froma called indel using bcftools (Li, 2011). Following read alignment andSNP calling and filtering, all mapping statistics and calculations weredone using R (RTeam, 2015). Read depth for each allele at segregatingbi-allelic SNPs in 1 Mb sliding windows (by 100 kb) was summed for thevarious mutant (lin, s2,j2, suppression of s2) and wild-type sequencingpools and mutant:non-mutant SNP ratios were calculated. Finally, mutantSNP ratio was divided by wild-type SNP ratio (+0.5) and plotted acrossthe tomato genome.

Data and Software Availability

Raw sequencing reads generated in this study have been deposited at theSequence Read Archive (ncbi.nlm.nih.gov/sra) under BioProject SRP100435.

Additional resources for the tomato core collection (please see e.g.,unity.phenome-networks.com), for CRISPR design (please see e.g.,cbi.hzau.edu.cn/cgi-bin/CRISPR), for sequence retrieval (please seee.g., phytozome.jgi.doe.gov/) and for data deposition (please see e.g.,ncbi.nlm.nih.gov/sra) are also available to one of ordinary skill in theart.

Results

The s2 Variants Produce Branched Inflorescences and Flowers withJointless Pedicels

To explore the challenges with improving tomato inflorescences, a corecollection of 4,193 wild and domesticated accessions was screened fordeviation from the typical inflorescence architecture of multipleflowers arranged along a single branch (FIG. 1A)(unity.phenome-networks.com, see STAR Methods). Twenty-three extremelybranched accessions were previously reported that were all defective inthe gene COMPOUND INFLORESCENCE (S, homolog of ArabidopsisWUSCHEL-RELATED HOMEOBOX 9, WOX9) (FIG. 1B) (Lippman et al., 2008).However, three rare variants not allelic to s that branched lessfrequently and also lacked the abscission zone on the stems (pedicels)of flowers known as the “joint” were also found (FIGS. 1C, 1D, and7A-7F). Searching other germplasm sources provided one additionalbranched jointless mutant derived from an X-ray mutagenesis (FIGS. 7Cand 7F) (Stubbe, 1972). Crosses among all four accessions failed tocomplement (FIGS. 7G-71). Consequently, these accessions werecollectively named compound inflorescence 2 (s2).

One s2 accession was designated as a reference (LA4371), and an analysisof higher-order mutants with s showed an additive genetic relationship,indicating the gene(s) underlying s2 function separately from the S gene(FIGS. 1C and 7J). It was noted during the generation of s s2 plantsthat s2 segregated at a ratio of ˜ 1/16 (FIG. 1E), suggesting twounlinked recessive mutations underlie s2 phenotypes. Consistent withthis, jointless plants (unbranched and branched) segregated as a singlerecessive mutation. This jointless trait resembled twoclassicaljointless-2 (j2) mutants reported 50 years ago. The original j2was discovered in the unbranched wild tomato species S. cheesmaniae fromthe Galapagos Islands (Rick, 1956a). A second allele arose spontaneouslyin an agricultural field, but this mutation was also associated withinflorescence branching that caused excessive flower production and poorfruit set due to epistatic interactions with the domesticated germplasm(Reynard, 1961; Rick, 1956b). Breeders selected and utilized unbranchedj2, because it reduced fruit dropping and enabled large-scale machineharvesting of processing tomatoes, while maintaining good fruit set(Robinson, 1980; Zahara and Scheuerman, 1988). Notably, the jointlessphenotype of s2 was allelic to j2 (FIG. 7K), and s2 plants with normalpedicels were not found, suggesting branching required the j2 mutation.Therefore, the second locus was designated enhancer-of-jointless2 (ej2).

To better understand the developmental basis of s2 branching, the stagesof meristem maturation during early inflorescence development wereexamined. Tomato inflorescences develop according to the sympodialgrowth program (Park et al., 2014a), in which each vegetative meristemmatures into a transition meristem (TM) and terminates in a floralmeristem (FM) that produces the first flower of the inflorescence.Additional flowers arise from iterative formation of specializedaxillary (sympodial) inflorescence meristems (SIM), resulting in amulti-flowered inflorescence (FIG. 1F). In s mutants, both TM and SIMmaturation are severely delayed, allowing multiple SIMs to form at eachcycle (FIG. 1G) (Lippman et al., 2008; Park et al., 2012). AdditionalSIMs also formed in s2 plants, but less than in s (FIG. 1H). Todetermine if s2 was delayed in maturation, RNA-seq was performed onsequential s2 meristem maturation stages and compared transcriptomedynamics with existing maturation profiles for s and WT (see STARMethods) (Park et al., 2012). A principal component analysis (PCA) using2,582 maturation marker genes (Lemmon et al., 2016) showed fewer TM andFM marker genes were delayed during s2 meristem maturation compared tos, consistent with less branching in s2 inflorescences (FIGS. 1I-1K and8).

Mutations in Two related MADS-Box Genes Cause s2 Branching

The j2 mutant was previously mapped to the centromere of chromosome 12,but poor recombination prevented identification of the responsible gene(Budiman et al., 2004; Yang et al., 2005). To clone the genes underlyingj2 and ej2, two F₂ populations were generated from crossing s2 with thejointed (J2/J2) cultivar M82 and the wild ancestor of tomato, S.pimpinellifolium. In the intra-species F₂ population, s2 plantssegregated at the expected ratio of ˜ 1/16, but this segregation wassubstantially lower in the S. pimpinellifolium population, suggestingunknown modifier loci can suppress s2 branching (FIGS. 9A-9C). To map j2and ej2 simultaneously, genome sequencing was performed on pools of DNAfrom s2,j2, and WT F₂ segregating plants (see STAR Methods). ComparingSNP ratios between s2 and WT pools in both populations revealed a regionnear the bottom of chromosome 3 and the centromere of chromosome 12 witha strong bias for SNPs from the s2 parent (FIGS. 2A, 9D, and 9E). SNPratios between s2 and j2 revealed a bias only at the bottom ofchromosome 3. These results confirmed j2 is located near the chromosome12 centromere and revealed ej2 resides on chromosome 3.

MADS-box transcription factors are known to contribute to pedicelabscission zone development in tomato (Liu et al., 2014; Mao et al.,2000; Nakano et al., 2012; Shalit et al., 2009). The jointless1 mutant(j1) was mapped to chromosome 11 and found to be mutated in a homolog ofthe Arabidopsis MADS-box flowering regulator SHORT VEGETATIVE PHASE(SVP) (Hartmann et al., 2000; Mao et al., 2000). Therefore, the ˜6 Mbpj2 mapping interval for MADS-box genes was searched, and among the 164genes in this region only one candidate was found, Solyc12g038510, ahomolog of the Arabidopsis floral organ identity MADS-box geneSEPALLATA4 (SEP4) (FIG. 2B) (Ditta et al., 2004). Previoustranscriptional silencing of Solyc12g038510 resulted in jointlesspedicels, but it was suggested Solyc12g038510 and J2 were differentgenes, because the published j2 mapping interval did not coincide withSolyc12g038510, likely from unreliable centromeric marker resolution(Budiman et al., 2004; Liu et al., 2014). However, the genomicsequencing of s2 and j2 mutants exposed a Copia/Rider-type transposableelement (TE) in the first intron of Solyc12g038510 that was absent in WT(FIG. 2C). Furthermore, the s2 RNA-seq revealed that most Solyc12g038510transcripts initiated in the first intron, resulting in an earlynonsense mutation (FIGS. 2D and 9H). To validate that Solyc12g038510 isJ2, CRISPR/Cas9 was used to engineer loss-of-function mutations, and theresulting j2^(CR) plants developed jointless unbranched inflorescences(FIGS. 2E and 2F). Moreover, progeny from crossing j2^(CR) withs2-derived j2 had jointless and unbranched inflorescences (FIG. 2G), andsequencing Solyc12g038510 in the original j2 S. cheesmaniae accessionrevealed an early stop codon (FIGS. 9F-9H). Thus, the SEP4 geneSolyc12g038510 is J2, and two natural mutations arose independently(hereafter designated j2^(TE) and j2^(stop)) (Reynard, 1961; Rick,1956a).

Both j2 and ej2 are required for s2 branching, suggesting the underlyinggenes function redundantly, similar to SEP genes in Arabidopsis thatcontrol floral organ identity (Ditta et al., 2004; Pelaz et al., 2000).The 66 genes were searched in the 500 kbp ej2 mapping interval forMADS-box genes and the tandemly arranged Solyc03g114830 andSolyc03g114840 were found (FIG. 2H). Solyc03g114830 is a homolog ofArabidopsis FRUITFULL and transcriptional knockdown of this gene causesdefects in fruit ripening (Bemer et al., 2012; Wang et al., 2014). Thegenomic sequencing of s2 mutants did not reveal any Solyc03g114830coding or noncoding SNPs, or large indels, and s2 fruits ripenednormally. In contrast, Solyc03g114840 is another homolog of SEP4, and a564 bp insertion was found in the 5^(th) intron of s2 mutants, which wasabsent in WT (FIG. 2I). Notably, RNA-seq reads from s2 revealed a thirdof Solyc03g114840 transcripts were misspliced, suggesting the insertioncaused a partial loss of function (FIG. 2J). To test this and uncoverthe phenotypic consequences of strong loss of EJ2 function, new alleleswere engineered with CRISPR/Cas9 and e j2^(CR) inflorescences were foundto be unbranched, but the sepals (outermost leaf-like organs of theflowers) were exceptionally large and fruits were pear-shaped (FIGS. 2Kand 2L). To determine if the original ej2 mutation impacted flowerand/or fruit morphology, ej2 was backcrossed into M82 and relative sepallength (defined by sepal/petal length ratio) was measured. Notably,whereas there was no obvious change in fruit shape or size, ej2 sepalswere 50% longer than WT but shorter than ej2^(CR), consistent with aweak allele (FIGS. 2M, 2N, and 9I). Importantly, flowers of F₁ progenyfrom crossing ej2 and ej2^(CR) also developed long sepals. Thus,Solyc03g114840 is E12, and the natural ej2 mutation is a weakloss-of-function allele (hereafter designated ej2^(w)).

Finally, it was verified that the other s2 accessions carried mutationsin both j2 and ej2. PCR genotyping showed all but one accession wasdouble mutant for ej2^(w) and either j2^(TE) or j2^(stop) (FIG. 9J). Thelast accession was homozygous for ej2^(w), but J2 could not beamplified, consistent with having originated from an X-ray mutagenesis(Stubbe, 1972). Thus, the prolonged meristem maturation underlying s2inflorescence branching is caused by mutations in two redundantly actingSEP MADS-box genes.

Ej2^(w) Arose During Domestication and Hindered j2 Utilization forBreeding

In modern breeding programs, the value of jointless varieties wasrecognized for their potential to reduce fruit drop and post-harvestdamage during mechanical harvesting for the processing tomato industry.Yet, plants carrying j1 yield poorly due to reversion of inflorescencesto vegetative growth after developing a few flowers (Butler, 1936; Maoet al., 2000). Thus, j2 was widely favored over the last 50 years ofbreeding. However, breeders frequently experienced problems withexcessive inflorescence branching and low yield upon introducing j2 intodifferent cultivated backgrounds (Robinson, 1980), probably because ofnegative epistasis with ej2^(w). To determine to what extent ej2^(w)hindered j2 utilization in breeding, 568 wild and domesticatedaccessions were genotyped from the tomato core collection and more thanhalf were found to be homozygous for the ej2^(w) allele (FIG. 3A).Notably, ej2^(w) was absent from S. pimpinellifolium, but 40% of earlydomesticates (landraces) were homozygous for the mutation, and thepercentage doubled in cultivars. Most importantly, ej2^(w) was stronglyassociated with long sepals, including within a subset of confirmedlandraces (Blanca et al., 2015), suggesting selection duringdomestication (FIGS. 3B-3E). In support of this, ej2^(w) is in closeproximity (<46 Kbp) to a previously reported domestication andimprovement selective sweep (Lin et al., 2014). Notably, a minor fruitweight QTL (fv3.2) that also arose in the landraces is in closeproximity (˜85 Kbp) to EJ2 (Chakrabarti et al., 2013; Zhang et al.,2012). Among 62 landraces, accessions were found that carried ej2^(w)but not fw3.2 (ej2^(w)/FW3.2: 7%) and vice versa (EJ2/fv3.2: 9%),suggesting that each mutant allele arose independently and were likelycombined early in domestication. It was also found that not allcultivated lines carried both alleles (ej2^(w)/FW3.2: 2%; EJ2/fv3.2:11%), indicating that both mutations were either passed on independentlyduring domestication and improvement, or were co-selected and thenseparated later by breeding.

One explanation for the early selection of ej2^(w) and its subsequentspread in the cultivated germplasm is that larger sepals provided anenlarged calyx that was concomitantly selected as fruit size increased,perhaps with fw3.2. Such a trait would not necessarily have beenselected for improved productivity by increasing fruit size or numberper se, but instead could have provided improved fruit support, stronglocal source tissue, or simply aesthetic value for larger fruits. Todetermine if ej2^(w) was selected during domestication and breeding oflarger fruits, the frequency of the ej2^(w) allele was evaluated in 258cultivars representing five fruit sizes ranging from small “cherry”tomatoes (<5 g) to extremely large “beefsteak” varieties (>500 g).Remarkably, the frequency of the allele increased with fruit size, andnearly all (>90%) large-fruited accessions were homozygous for ej2^(w),including 88% of vintage heirloom cultivars (Male, 1999). These resultsshow that the ej2^(w) allele was already widespread in larger fruittypes before j2 was discovered and adopted in modern breeding (FIG. 3F).Since EJ2 is also expressed in developing fruits (FIG. 10A) and ej2^(CR) fruits are elongated (FIG. 2L), it is also possible the ej2^(w)allele impacts other fruit traits such as size/shape and/or ripening,especially in the presence of other QTL that impact these traits.

Elite breeding germplasm carries both j2^(TE) and ej2^(w), but branchingis suppressed

Because ej2^(w) became widespread in tomato germplasm and j2 arose muchlater, introducing either of the j2 alleles into most cultivars wouldhave resulted in undesirable branching and low yield. However, it wasreported these adverse effects could be overcome by breeding (Robinson,1980). One possibility is that ej2^(w) was segregated away throughcrosses. Alternatively, breeders could have identified and selectednatural suppressors of branching. To test this, 153 unbranched jointedand jointless elite inbreds and hybrids were obtained from major seedcompanies and public breeders (see STAR Methods), and genotyped for bothmutations. All jointless lines were homozygous for j2^(TE), indicatingthe allele that arose in the domesticated germplasm was favored inbreeding. Since new tomato varieties for processing and fresh-marketproduction are developed in separate breeding programs, it was asked ifj2^(TE) was utilized in both. The value of the jointless trait is mostrecognized for mechanical harvesting of processing types, and in supportof this the j2^(TE) allele was present in 74% of sampled processinglines. Although less widespread, j2^(TE) was also found in 34% offresh-market lines, indicating that j2^(TE) continues to be utilized inboth breeding programs.

Remarkably, it was found that more than 60% of j2^(TE) homozygotes inboth processing and fresh-market lines were also homozygous ej2^(w)(FIGS. 4A and 4B), supporting the hypothesis that suppressors wereselected during improvement. This was reminiscent of the reducedsegregation of s2 in the S. pimpinellifolium F₂ mapping population(FIGS. 9B and 9C). To map potential suppressor loci, 1,536 F₂ plantswere regrown, and of 92 plants homozygous for both mutations, 24% showedvarious degrees of suppression (FIG. 4C).

Using genome sequencing, one large-effect suppressor was mapped near theend of chromosome 2 in the same region as a previously reportedsuppressor in the domesticated germplasm (FIG. 4D) (Robinson, 1980).However, given that only a small percentage of j2^(TE) ej2^(w) F₂ plantsdisplayed unbranched inflorescences, additional suppressors frombreeding germplasm are likely involved, which together were needed toachieve complete suppression.

Three meristem expressed SEP4 genes modulate inflorescence complexity

The dissection of the negative epistasis underlying s2 branching exposedtwo tomato SEP4 genes that act redundantly to control meristemmaturation and inflorescence development. This led to the question of towhat extent these genes work with other tomato SEP family members toregulate inflorescence architecture and flower production, and couldhave potential for agricultural application. In Arabidopsis, a family offour redundant SEP genes is required to establish floral organ identity(Ditta et al., 2004; Pelaz et al., 2000). Tomato has an expanded SEPfamily of six members (Consortium, 2012), and a phylogenetic analysis ofprotein sequences showed Arabidopsis SEP1, 2, and 3 have two tomatohomologs (Solyc05g015750/TM5 and Solyc02g089200/TM29) (FIG. 5A). Incontrast, there are four homologs of SEP4, and among them is theRIPENING INHIBITOR (RIN) gene. A classical mutation in RIN blocksripening and is widely used in hybrid breeding due to a heterozygousdosage effect that causes fruits to remain firm and ripen over aprotracted period, improving shelf life (Klee and Giovannoni, 2011;Vrebalov et al., 2002).

To investigate individual and combined roles of tomato SEP genes ininflorescence development, expression patterns were first analyzed usingthe meristem maturation atlas and transcriptome data from other majortissues (Consortium, 2012; Park et al., 2012). Both TM5 and TM29(SEP1/2/3 homologs) were expressed only later in reproductivedevelopment, beginning in floral meristems and extending into flowersand fruits (FIGS. 5B and 10A), supporting previously characterized rolesin floral organ identity (Ampomah-Dwamena et al., 2002; Pnueli et al.,1994). RIN was only expressed in fruits, consistent with its role inripening (FIG. 10A) (Vrebalov et al., 2002). In contrast, expression ofJ2, EJ2, and the fourth SEP4 homolog (Solyc04g005320) began earlier, inthe TM stage of meristem maturation and in SIMs (FIG. 5B). Thissuggested Solyc04g005320 could function with J2 and EJ2 in meristemmaturation. Moreover, given that Arabidopsis SEP redundancy is based onformation of multimeric protein complexes (Theissen et al., 2016),interactions were tested among all four tomato SEP4 proteins in yeasttwo-hybrid assays and J2, EJ2, and Solyc04g005320 were found to interactwith each other and themselves, except for homomeric EJ2. These resultsvalidated previous findings (Leseberg et al., 2008), and furtherrevealed that J2 and EJ2 interact with each other, supporting redundancyin the control of meristem maturation and inflorescence architecture(FIGS. 5C, 5D, 10B and 10C).

To test if Solyc04g005320 contributes to inflorescence architecture andflower production, CRISPR/Cas9 was used to engineer plants with nullmutations, which resulted in exceptionally long inflorescences withnearly twice as many flowers as WT and longer internodes (FIGS. 5E and10D). Weak branching late in inflorescence development was alsofrequently observed. Whether similar effects occur in genotypes thatalready have long inflorescences was tested by mutating Solyc04g005320in S. pimpinellifolium, which produces 15-20 flowers on eachinflorescence. Remarkably, internode length and flower number doubled(FIGS. 5F, 10D-10F). These phenotypes were reminiscent of agamma-irradiation mutant designated long inflorescence (lin) that waspreviously mapped to an interval on chromosome 4 containingSolyc04g005320 (FIGS. 10G-10J)(see STAR Methods). SequencingSolyc04g005320 from the lin mutant revealed a translocation in the firstintron that eliminated transcription (FIGS. 10J-10L, also referred toherein as lin^(trans)) and crosses with a CRISPR allele failed tocomplement the long inflorescence phenotype.

The increase in inflorescence complexity in lin mutants is modestcompared to j2 ej2^(w) double mutants. To study the extent of redundancyand potential dosage relationships among the three genes, strong alleleswere used in the same background to create all combinations ofhigher-order mutants (see STAR Methods). Whereas j2^(CR) was largelyadditive with lin (FIG. 10M), ej2^(CR) and lin were synergistic forfloral organ development; double-mutants had long inflorescences withmore flowers that developed extremely enlarged sepals, but inner floralorgans did not fully develop and fruits failed to form (FIG. 10N). Asexpected, j2^(CR) and ej2^(CR) were also synergistic, but unlike themoderately branched, highly floral inflorescences of the originalj2^(TE/stop) ej2^(w) natural double mutants (s2), inflorescences fromj2^(CR) e j2^(CR) plants were extraordinarily branched and rarelyproduced normal fertile flowers (FIG. 5G). Finally, combining all threemutants resulted in massively over proliferated SIMs without formingflowers (FIGS. 5H and 10O). The same effect was observed in S.pimpinellifolium j2^(CR) ej2^(CR) lin^(CR) plants (FIGS. 5I and 10O).The sequences for S. pimpinellifolium j2^(CR) ej2^(CR) lin^(CR) areshown below. Thus, J2 and E/2 have distinct roles in floral development,but all three SEP4 genes have overlapping roles in meristem maturationand inflorescence development.

Dosage of Meristem Maturation Transcription factors can be Exploited toImprove Inflorescence Architecture and Yield

The individual and combined mutations in J2, EJ2, and LIN provided aseries of three forms of increased inflorescence complexity ranging fromweak (lin single mutants) to extremely severe (j2 ej2 lin triplemutants), indicating quantitative relationships among these SEP4 genes.It was previously demonstrated that dosage relationships among genes inthe florigen pathway could be exploited to create a quantitative rangeof plant architectures that translated to improved productivity indeterminate field-grown tomatoes (Park et al., 2014b; Soyk et al.,2016). It was reasoned that dosage sensitivity could be similarly usedto fine-tune inflorescence architecture and flower production. To testthis, a series of homozygous and heterozygous combinations of j2 strongalleles with ej2^(w) or ej2^(CR) in the isogenic M82 background wasfirst created (FIGS. 6A and 6B). All double heterozygotes (e.g.j2/+ej2^(w)/+; j2/+ej2^(CR)/+) and plants heterozygous for j2 andhomozygous for ej2^(w) (j2/+ej2^(w)) produced unbranched inflorescenceslike the single mutants. In contrast, heterozygosity for ej2^(w) in a j2background (j2 ej2^(w)/+) conferred weak branching, as did j2/+ej2^(CR).Notably, heterozygosity for the null e j2^(CR) allele in the null j2background (j2 ej2^(CR)/+) resulted in branching that matched s2inflorescences (j2 ej2^(w)), further validating that ej2^(w) is a weakallele and confirming a sensitive dosage relationship between thesegenes. Given these results, it was reasoned that other meristemmaturation regulators might have similar dosage-sensitivity oninflorescence architecture and this was tested with S, a member of theWOX protein family (Graaff et al., 2009; Lippman et al., 2008). Indeed,plants heterozygous for three s mutant alleles were also mildly branched(FIGS. 6C and 6D), demonstrating dosage-sensitivity of independentmeristem maturation genes allows for quantitative tuning ofinflorescence architecture.

DISCUSSION Dose-Dependent Quantitative Variation, Weak Alleles, and CropImprovement

This study involved exploration of the potential of genes and allelesunderlying natural variation in inflorescence complexity to improveproductivity. By analyzing the s2 branching variant, it was found thatmultiple members of the SEP4 subfamily of tomato MADS-box genes playcritical redundant roles in modulating meristem maturation andinflorescence architecture. The first MADS-box family member involved intomato domestication was further described, highlighting the growingsignificance of this transcription factor family in contributing todomestication and improvement of diverse crops (Singh et al., 2013;Vrebalov et al., 2002; Zhao et al., 2011). By dissecting interactionsbetween meristem expressed SEP4 genes dosage relationships wereuncovered among an allelic series of MADS-box mutations with potentialfor breeding. This collection of alleles, including mutations in S,comprises a toolkit to manipulate inflorescence architecture, which cannow be expanded to additional regulators of meristem maturation, such asLIN. To demonstrate this, CRISPR/Cas9 was used to target LIN in theelite cherry tomato cultivar Sweet 100 and mutant lines were generatedwith moderately branched inflorescences and increased flower production(FIGS. 10P-10S).

The present approach for creating desirable phenotypic variation inmajor yield traits relies on combining specific heterozygous andhomozygous mutations to obtain a quantitative range of dosage effects(Park et al., 2014b). However, exploiting gene dosage may be limited bythe availability of weak alleles that confer quantitative traitmodifications. For example, longer sepals and weak branching wereachieved through different levels of reduced EJ2 dosage fromhomozygosity and heterozygosity for ej2′, respectively. In nature,similar dosage effects often arise from mutations in transcriptionalcontrol regions (e.g., in cis-regulatory DNA). Such alleles were widelyfavored in crop domestication and improvement for their subtlephenotypic changes compared to null alleles that frequently displaydeleterious pleiotropic effects (Meyer and Purugganan, 2013; Puruggananand Fuller, 2009). For example, increased fruit size during tomatodomestication depended in part on transcriptional alleles of multiplecomponents in the classical CLAVATA-WUSCHEL stem cell circuit (Xu etal., 2015). A potentially powerful approach to engineer novel weakalleles that are being explored (Swinnen et al., 2016) is exploitinggene-editing technology to mutate cis-regulatory control regions ofproductivity genes. A promising target identified in this study is LIN.CRISPR/Cas9-induced weak transcriptional alleles that confer reduced LINexpression may provide subtle increases in flower production, which maybe especially valuable in large-fruited cultivars where branching oftennegatively impacts fruit weight and yield. Notably, a rice homologofLINand other meristem maturation genes control panicle architectureand grain production (Kobayashi et al., 2010, 2012; Liu et al., 2013),suggesting the present findings have broad agricultural potential. Newgene-editing tools should enable the engineering of diverse types andstrengths of alleles that can provide customized gene dosage effects toimprove a wide range of agronomic traits in many crops.

Epistasis in Evolution, Domestication, and Breeding

Progress in breeding is largely driven by loci with predictable additiveeffects. For example, the majority of flowering time variation in maizeis determined by thousands of small additive quantitative trait loci(QTL) (Buckler et al., 2009), and the same is true for traits in othercrops (Doust et al., 2014; Gao et al., 2015). Yet, epistaticinteractions, both positive and negative, are also important inbreeding, particularly when working with disparate germplasm. Forexample, interactions between interspecific quantitative trait loci(QTL) in rice can improve aluminum tolerance (Famoso et al., 2011),whereas stacking multiple wild species-derived QTL affecting the sameyield traits in tomato results in less-than-additive or “diminishingreturns” epistasis (Eshed and Zamir, 1996).

In recent years, several cases of negative epistasis have emerged indiverse organisms involving clashes between newly evolved andestablished alleles, or upon bringing together distinct genomes, eitherthrough natural or artificial means. Examples include compromisedfitness gains upon combining interacting alleles in bacteria and yeast(Chou et al., 2011; Heck et al., 2006; Khan et al., 2011; Kvitek andSherlock, 2011), hybrid necrosis between distinct accessions ofArabidopsis (Chae et al., 2014), and loss of protection from malaria inhumans when two common resistance variants are co-inherited (Williams etal., 2005). Compared to negative epistasis in evolution and naturalselection, the intense artificial selection imposed by humans duringdomestication and breeding could drive more frequent occurrences ofepistasis. While dramatic cases like the one described in this studycould be overcome through selection against interactions or suppressionwith modifiers, there may be many undiscovered negative interactions inagriculture with more subtle phenotypic consequences that may remainchallenging to detect and dissect until high throughput quantitativephenotyping platforms (phenomics) and power in genome-wide associationstudies (GWAS) improves.

The present dissection of the extreme negative epistasis underlying thes2 branching syndrome has highlighted an underappreciated challenge forthe next generation of crop breeding. Specifically, using rapidlyadvancing gene-editing technologies to introduce precise novel allelicvariation for specific genes into existing germplasm may not providedesirable phenotypic outcomes, and could potentially result in negativeconsequences due to interactions with alleles selected and stabilizedduring domestication and early breeding (Mackay, 2013). That the presentexample of negative epistasis involved two closely related MADS-boxgenes suggests that engineering new alleles in gene families or relateddevelopmental pathways that already played a role in domestication andimprovement may be particularly sensitive to unexpected epistaticconsequences, perhaps explaining other as yet uncharacterized examplesof negative epistasis in agriculture (Bomblies and Weigel, 2007;Matsubara et al., 2015; Shang et al., 2016; Zhang et al., 2011).Elucidating, neutralizing, and potentially exploiting negativeepistasis, as done in the present study, could have a significant impactin helping break productivity barriers in breeding of both plants andanimals.

TABLE 2 Oligos used in this study Yeast two-hybrid assays Gene ForwardReverse Restr. name Gene ID primer primer enzyme LIN Solyc04 CACCGAATTCATTCGGATCCTCA EcoRI + g005320 TGGGAAGAGGT AAGCATCCATCC BamHI AAGGTAGAATGGTAA (SEQ (SEQ ID NO: ID NO: 18) 17) J2 Solyc12 CACCGAATTCATTCGGATCCTTA EcoRI + g038510 TGGGAAGAGGA GAGCATCCACCC BamH I AGAGTAGAACTGGAAT (SEQ (SEQ ID NO: ID NO: 20) 19) EJ2 Solyc03 CACCGAATTCATTCGGATCCTTA EcoRI + g114840 TGGGAAGAGGA AAGCATCCATCC BamHI AGAGTTGAGATGAATAAATC (SEQ ID NO: (SEQ ID NO: 21) 22) RIN Solyc05 CACCGAATTCATTCGGATCCTCA EcoRI + g012020 TGGGTAGAGGG AAGCATCCATCC BamH I AAAGTAGAAAGGTACA (SEQ (SEQ ID NO: ID NO: 24) 23)Natural and induced mutant alleles analyzed in this study, and respectivegenotyping markers Gene Forward Reverse Allele Restr. name Gene IDprimer primer name Type WT Mutant enzyme LIN Solyc04 GCAAAACTTTACTTTTTGATTCA lin indel 396 — — g005320 AATTAGTTCTA TGTGTCTGTAC ATG (SEQ(SEQ ID NO: ID NO: 25) 26) LIN Solyc04 AATATCGTGTT CTTTTTGATTCA linindel —  358 — g005320 AGAATGTGACA TGTGTCTGTAC C (SEQ ID (SEQ ID NO:NO: 27) 28) J2 Solyc12 TTACTTTTGCT CCGTCCTTTCTG j2-TE Indel —  193 —g038510 AAGAGAAGAAA TTTGTAGC bp TGG (SEQ ID (SEQ ID NO: NO: 29) 30) J2Solyc12 TTACTTTTGCT GAATCCACTTAA j2-TE Indel 709 — g038510 AAGAGAAGAAAGAATCTCTACC bp TGG (SEQ ID (SEQ ID NO: NO: 31) 32) J2 Solyc12TATTGTGATAT AATACCTGAGTA j2- dCAPS 206  230 HpaI g038510 GTAGAGTGGTGTCACTAACCGTT classic bp + bp C (SEQ ID (SEQ ID NO: 24 NO: 33) 34) bp EJ2Solyc03 CACAATTCATG CGGAGTAATCTA ej2-w Indel 177  738 — g114840CTGGATCAGC TTAGATTCTGC bp bp (SEQ ID NO: (SEQ ID NO: 35) 36) LIN Solyc04CCTTTAATAAG TTGAAGGTGCAT CR-lin- Indel 855 1390 — g005320 TTGAAAATCCCAGAACATACC a1 bp bp TC (SEQ ID (SEQ ID NO: NO: 37) 38) LIN Solyc04CCTTTAATAAG TTGAAGGTGCAT CR-lin- CAPS 796  855 Hinc g005320 TTGAAAATCCCAGAACATACC a2 bp  bp II TC (SEQ ID (SEQ ID NO:  59 NO: 39) 40) bp J2Solyc12 ATATTGAATCG TAACTTTCTTCA CR-j2- Indel 316  411 — g038510TGTGATTGTCT AAGATGCATCC a1 bp bp C (SEQ ID (SEQ ID NO: NO: 41) 42) J2Solyc12 ATATTGAATCG TAACTTTCTTCA CR-j2- CAPS 316  178 MboI g038510TGTGATTGTCT AAGATGCATCC a2 bp bp  I C (SEQ ID (SEQ ID NO:  139 NO: 43)44) bp EJ2 Solyc03 AATATGGTCCT TAGCAAACTTAT CR-ej2- Indel 236  211 —g114840 TATGTCCAACC TGGGCTAGC a1 bp bp (SEQ ID NO: (SEQ ID NO: 45) 46)EJ2 Solyc03 AATATGGTCCT TAGCAAACTTAT CR-ej2- CAPS 236  144 Hind g114840TATGTCCAACC TGGGCTAGC a2 bp bp  III (SEQ ID NO: (SEQ ID NO:   94 47) 48)bp Cas9 — CTGACGTAAGG CATCTCATTACT — T- —  446 — GATGACGCAC AAAGATCTCCDNA bp (SEQ ID NO: (SEQ ID NO: 49) 50) RT-PCR Gene Forward Reverse nameGene ID primer primer LIN Solyc04 ATGGGAAGAGG TCAAAGCATCCA g005320TAAGGTAGAA TCCTGGTAAA (SEQ ID NO: (SEQ ID NO: 51) 52) J2 Solyc12ATGGGAAGAGG TTAGAGCATCCA g038510 AAGAGTAGAAC CCCTGGAAT (SEQ ID NO:(SEQ ID NO: 53) 54) EJ2 Solyc03 ATGGGAAGAGG TTAAAGCATCCA g114840AAGAGTTGAG TCCATGAATAAA (SEQ ID NO: TC (SEQ ID 55) NO: 56) UBI Solyc01CGTGGTGGTGC ACGAAGCCTCTG g056940 TAAGAAGAG AACCTTTC (SEQ ID NO:(SEQ ID NO: 57) 58) CRISPR/Cas9 genome-editing sgRNA Forward ReversesgRNA name Gene ID primer primer sequence LIN- Solyc04 TGTGGTCTCAATGTGGTCTCAAG TTCTAGT sgRNA- g005320 TTTTCTAGTAT CGTAATGCCAAC ATGTCTG 1GTCTGATACAC TTTGTAC (SEQ ATACAC GTTTTAGAGCT ID NO: 60) (SEQ IDAGAAATAGCAA NO: 81) G (SEQ ID NO: 59) LIN- Solyc04 TGTGGTCTCAATGTGGTCTCAAG GGAACAG sgRNA- g005320 TTGGAACAGCT CGTAATGCCAAC CTTGAGC 2TGAGCGTCAAC TTTGTAC (SEQ GTCAAC GTTTTAGAGCT ID NO: 62) (SEQ IDAGAAATAGCAA NO: 82) G (SEQ ID NO: 61) J2- Solyc12 TGTGGTCTCAATGTGGTCTCAAG AGCTCCT sgRNA- g038510 TTAGCTCCTTC CGTAATGCCAAC TCAACGT 1AACGTTCTCAA TTTGTAC (SEQ TCTCAA GTTTTAGAGCT ID NO: 64) (SEQ IDAGAAATAGCAA NO: 83) G (SEQ ID NO: 63) J2- Solyc12 TGTGGTCTCAATGTGGTCTCAAG ACATATT sgRNA- g038510 TTACATATTCT CGTAATGCCAAC CTTGGAG 2TGGAGAGGATT TTTGTAC (SEQ AGGATT GTTTTAGAGCT ID NO: 66) (SEQ IDAGAAATAGCAA NO: 84) G (SEQ ID NO: 65) EJ2- Solyc03 TGTGGTCTCAATGTGGTCTCAAG TTTGGGC sgRNA- g114840 TTTTTGGGCAC CGTAATGCCAAC ACGTTAA 1GTTAAGCTCGA TTTGTAC (SEQ GCTCGA GTTTTAGAGCT ID NO: 68) (SEQ IDAGAAATAGCAA NO: 85) G (SEQ ID NO: 67) E32- Solyc03 TGTGGTCTCAATGTGGTCTCAAG CCTTAAA sgRNA- g114840 TTCCTTAAAGC CGTAATGCCAAC GCAAATC 2AAATCAGGTCA TTTGTAC (SEQ AGGTCA GTTTTAGAGCT ID NO: 70) (SEQ IDAGAAATAGCAA NO: 86) G (SEQ ID NO: 69) LIN/ Solyc12 TGTGGTCTCAATGTGGTCTCAAG GCTTTTG J2/ g038510; TTGCTTTTGCT CGTAATGCCAAC CTAAGAG EJ2-Solyc04 AAGAGAAGAAA TTTGTAC (SEQ AGAA sgRNA- g005320; GTTTTAGAGCTID NO: 72) (SEQ ID 1 Solyc03 AGAAATAGCAA NO: 87) g114840 G (SEQ IDNO: 71) LIN/ Solyc04 TGTGGTCTCAA TGTGGTCTCAAG GCAGTCT J2/ g005320TTGCAGTCTTC CGTAATGCCAAC TCAAAGG EJ2- AAAGGATTCAC TTTGTAC (SEQ ATTCACsgRNA- GTTTTAGAGCT ID NO: 74) (SEQ ID 2 AGAAATAGCAA NO: 88) G (SEQ IDNO: 73) Sequencing Forward Reverse Target primer primer pSC- GTAAAACGACGCAGGAAACAGCT B- GCCAG (SEQ ATGAC (SEQ amp/ ID NO: 75) ID NO: 76) kanpICH TCCTGTCAAAC TAATGTACTGGG 47761 ACTGATAG GTGGATGCAG (SEQ ID NO:(SEQ ID NO: 77) 78) pAGM ATAAGCCCATC CGGATAAACCTT 4723 AGGGAGCAGTTCACGCC (SEQ ID NO: (SEQ ID NO: 79) 80)

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From the above description, one skilled in the art can easily ascertainthe essential characteristics of the present disclosure, and withoutdeparting from the spirit and scope thereof, can make various changesand modifications of the disclosure to adapt it to various usages andconditions. Thus, other embodiments are also within the claims.

What is claimed is:
 1. A genetically-altered Solanaceae plant comprisinga mutant Solyc04g005320 gene or a homolog thereof.
 2. Thegenetically-altered Solanaceae plant of claim 1, wherein the mutantSolyc04g005320 gene or homolog thereof is a null allele or a hypomorphicallele.
 3. The genetically-altered Solanaceae plant of claim 1, whereinthe genetically-altered Solanaceae plant is heterozygous or homozygousfor the mutant Solyc04g005320 gene or homolog thereof.
 4. Thegenetically-altered Solanaceae plant of claim 1, wherein thegenetically-altered Solanaceae plant further comprises a mutantSolyc12g038510 gene or a homolog thereof, a mutant Solyc03g114840 geneor a homolog thereof, or both a mutant Solyc12g038510 gene or a homologthereof and a mutant Solyc03g114840 gene or a homolog thereof.
 5. Thegenetically-altered Solanaceae plant of claim 1, wherein the plantfurther comprises a mutant Solyc12g038510 gene or homolog thereof andthe mutant Solyc12g038510 gene or homolog thereof is a null allele or ahypomorphic allele.
 6. The genetically-altered Solanaceae plant of claim5, wherein the genetically-altered Solanaceae plant is heterozygous orhomozygous for the mutant Solyc12g038510 gene or homolog thereof.
 7. Thegenetically-altered Solanaceae plant of claim 1, wherein the plantfurther comprises a mutant Solyc03g114840 gene or a homolog thereof andthe mutant Solyc03g114840 gene or homolog thereof is a null allele or ahypomorphic allele.
 8. The genetically-altered Solanaceae plant of claim7, wherein the genetically-altered Solanaceae plant is heterozygous orhomozygous for the mutant Solyc03g114840 gene or homolog thereof.
 9. Thegenetically-altered Solanaceae plant of claim 1, wherein the plantfurther comprises both a mutant Solyc12g038510 gene or a homolog thereofand a mutant Solyc03g114840 gene or a homolog thereof, each of which areindependently a null allele or a hypomorphic allele.
 10. Thegenetically-altered Solanaceae plant of claim 9, wherein thegenetically-altered Solanaceae plant is heterozygous or homozygous forthe mutant Solyc12g038510 gene or homolog thereof and is heterozygous orhomozygous for the mutant Solyc03g114840 gene or homolog thereof. 11.The genetically-altered Solanaceae plant of claim 1, wherein thegenetically-altered Solanaceae plant further comprises a mutantSolyc12g038510 gene or homolog thereof, and a mutant Solyc03g114840 geneor homolog thereof, and wherein each is a hypomorphic allele.
 12. Agenetically-altered Solanaceae plant, comprising a mutant Solyc12g038510gene or a homolog thereof and a mutant Solyc03g114840 gene or a homologthereof, wherein the genetically-altered Solanaceae plant is homozygousfor the mutant Solyc12g038510 gene or homolog thereof and heterozygousfor the mutant Solyc03g114840 gene or homolog thereof.
 13. Thegenetically-altered Solanaceae plant of claim 12, wherein the mutantSolyc12g038510 gene or homolog thereof is a null allele or a hypomorphicallele and the mutant Solyc03g114840 gene or homolog thereof is a nullallele or a hypomorphic allele.
 14. The genetically-altered Solanaceaeplant of claim 1, wherein the genetically-altered Solanaceae plant is atomato (Solanum lycopersicum) plant.
 15. The genetically-alteredSolanaceae plant of claim 4, wherein the mutant Solyc04g005320 gene orhomolog thereof, the mutant Solyc12g038510 gene or homolog thereof,and/or the mutant Solyc03g114840 gene or homolog thereof is introducedby technical means.
 16. The genetically-altered Solanaceae plant ofclaim 4, wherein the mutant Solyc04g005320 gene or homolog thereof, themutant Solyc12g038510 gene or homolog thereof, and/or the mutantSolyc03g114840 gene or homolog thereof is introduced by chemical orphysical means.
 17. The genetically-altered Solanaceae plant of claim16, wherein the mutant Solyc04g005320 gene or homolog thereof, themutant Solyc12g038510 gene or homolog thereof, and/or the mutantSolyc03g114840 gene or homolog thereof is introduced using CRISPR/Cas9,chemical mutagenesis, radiation, Agrobacterium-mediated recombination,viral-vector mediated recombination, or transposon mutagenesis. 18.(canceled)
 19. A crop harvested from genetically-altered Solanaceaeplants as defined in claim
 1. 20. A seed for producing agenetically-altered Solanaceae plant of claim
 1. 21. A method forproducing a genetically-altered Solanaceae plant, the method comprisingintroducing a mutation into a Solyc04g005320 gene or a homolog thereofin a Solanaceae plant, thereby producing a genetically-alteredSolanaceae plant containing a mutant Solyc04g005320 gene or homologthereof.
 22. The method of claim 21, wherein the mutation is introducedusing CRISPR/Cas9.
 23. The method of claim 21, wherein the mutationproduces a null allele or a hypomorphic allele of the Solyc04g005320gene or homolog thereof.
 24. The method of claim 21, wherein the methodfurther comprises introducing into the Solanaceae plant a mutation intoa Solyc12g038510 gene or a homolog thereof, introducing a mutation intoa Solyc03g114840 gene or a homolog thereof, or introducing the mutationinto the Solyc12g038510 gene or homolog thereof and introducing themutation into the Solyc03g114840 gene or homolog thereof.
 25. The methodof claim 24, wherein the mutation(s) is/are introduced usingCRISPR/Cas9.
 26. The method of claim 21, wherein the genetically-alteredSolanaceae plant containing the mutant Solyc04g005320 gene or homologthereof is crossed to another genetically-altered Solanaceae plantcomprising a mutant Solyc12g038510 gene or homolog thereof, a mutantSolyc03g114840 gene or homolog thereof, or both the mutantSolyc12g038510 gene or homolog thereof and the mutant Solyc03g114840gene or homolog thereof, thereby producing a genetically-alteredSolanaceae plant containing the mutant Solyc04g005320 gene or homologthereof and the mutant Solyc12g038510 gene or homolog thereof, themutant Solyc03g114840 gene or homolog thereof, or both the mutantSolyc12g038510 gene or homolog thereof and the mutant Solyc03g114840gene or homolog thereof.
 27. The method of claim 21, wherein thegenetically-altered Solanaceae plant is a tomato (Solanum lycopersicum)plant.
 28. A genetically-altered Solanaceae plant produced or obtainableby the method of claim
 21. 29. The genetically-altered Solanaceae plantof claim 1, wherein the mutant Solyc04g005320 gene or homolog thereof isa hypermorphic allele.
 30. The genetically-altered Solanaceae plant ofclaim 29, wherein the genetically-altered Solanaceae plant isheterozygous or homozygous for the mutant Solyc04g005320 gene or homologthereof.
 31. (canceled)